The Paleocene-Eocene Thermal Maximum (PETM; ca. 55.8 Ma) is thought to coincide with a profound but entirely transient change among nannoplankton communities throughout the ocean. Here we explore the ecology of nannoplankton during the PETM by using multivariate analyses of a global data set that is based upon the distribution of taxa in time and space. We use these results, coupled with stable isotope data and geochemical modeling, to reinterpret the ecology of key genera. The results of the multivariate analyses suggest that the community was perturbed significantly in coastal and high-latitudes sites compared to the open ocean, and the relative influence of temperature and nutrient availability on the assemblage varies regionally. The open ocean became more stratified and less productive during the PETM and the oligotrophic assemblage responded primarily to changes in nutrient availability. Alternatively, assemblages at the equator and in the Southern Ocean responded to temperature more than to nutrient reduction. In addition, the assemblage change at the PETM was not merely transient—there is evidence of adaptation and a long-term change in the nannoplankton community that persists after the PETM and results in the disappearance of a high-latitude assemblage. The long-term effect on communities caused by transient warming during the PETM has implications for modern-day climate change, suggesting similar permanent changes to nannoplankton community structure as the oceans warm.

Selected shell specimens of extant and fossil streamlined pectinids, which have or presumably had level-swimming ability, were examined experimentally to elucidate their hydrodynamic properties, in particular, airfoil efficiency estimated by lift-drag ratio. Using a stationary water tank for nautical engineering, lift and drag forces were measured at various attack angles. Of the examined species, Amusium japonicum, which is characterized by an unusually shiny surface, upward-cambered commissure and sharpened trailing edge, is the most efficient level swimmer and has the lowest drag coefficient and the highest value of lift-drag ratio at any attack angle. Amussiopecten praesignis from the Plio-Pleistocene may have swum horizontally because its airfoil efficiency is superior to that of a living level swimmer, Placopecten magellanicus. Although its shell shape is analogous to that of Placopecten, Camptonectes (Maclearnia) cinctus from the Lower Cretaceous shows a much inferior efficiency and a significant flow separation from the surface. Bernoulli's effect in convex-upward species may contribute to increase lift, but a certain attack angle is always required for level flight. The strategy of level swimming probably evolved independently in several pectinacean lineages in which swimming rather than shell robustness became the preferred defense against predators. One problem that must be solved is that some feedback mechanism is required to check pitching, rolling, and yawing of the shell to attain stability in level flight.

The evolutionary origin of Morozovella angulata from its immediate ancestor, Praemurica uncinata, is preserved in Paleocene sediments from the Gulf of Mexico. This event represents the beginning of the morozovellid radiation and marks the first appearance of keeled planktonic foraminifera after the Cretaceous/Tertiary extinction. Parallel biometric and isotopic analyses were performed on size-segregated specimens from a succession of stratigraphic horizons. The biometric data reveal a temporal pattern of variation consistent with paedomorphosis. The appearance of angulose juvenile chambers in the otherwise rounded ancestral form (Praemurica uncinata) results in an allometry that becomes more pronounced upsection. At the origin of M. angulata, the juvenile morphology of the ancestor is retained throughout the entire ontogeny. Isotopic analysis of this sequence reveals the gradual acquisition of an increasingly heavy adult δ13C signal relative to that of the juvenile, while the δ18O data display no temporal or size-related trends. The temporal increase seen in the slope of the δ13C/size relationship may reflect the evolution of an increased dependency on photosymbionts.

Interior chamber walls of ammonites range from smoothly undulating surfaces in some taxa to complex surfaces, corrugated on many scales, in others. The ammonite suture, which is the expression of the intersection of these walls on the exterior of the shell, has been used to assess anatomical complexity. We used the fractal dimension to measure sutural complexity and to investigate complexity over evolutionary time and showed that the range of variation in sutural complexity increased through time. In this paper we extend our analyses and consider two new parameters that measure the range of scales over which fractal geometry is a satisfactory metric of a suture. We use a principal components analysis of these parameters and the fractal dimension to establish a two-dimensional morphospace in which the shapes of sutures can be plotted and in which variations and evolution of suture morphology can be investigated. Our results show that morphospace coordinates of ammonitic sutures correspond to visually perceptible differences in suture shape. However, three main classes of sutures (goniatitic, ceratitic, and ammonitic) are not unambiguously discriminated in this morphospace. Interestingly, ammonitic sutures occupy a smaller morphospace than other suture types (roughly one-half of the morphospace of goniatitic and ceratitic sutures combined), and the space they occupied did not change dimensions from the Jurassic to the late Cretaceous.

We also compare two methods commonly used to measure the fractal dimension of linear features: the Box method and the Richardson (or divider) method. Both methods yield comparable results for ammonitic sutures but the Richardson method yields more precise results for less complex sutures.

Of all sessile filtrators, only some species of acorn barnacles managed to permanently settle on whales. Their key exaptation was probably a kind of biochemical cleaning process, which could be modified to penetrate into the host's dead cutis. Anchorage was further increased by coring prongs out of the whale skin (Coronula) or by transforming the wall into a cylindrical tube that added new rings at the base, while old ones flaked off at the surface in tandem with skin shedding (Tubicinella). Xenobalanus even everted its naked body into a stalked structure and reduced the wall plates to a minute, but highly efficient, anchor. Cryptolepas combines the strategies of Tubicinella and Coronula, but with a different structure of the radial folds. Because of a shared exaptational inventory, it is impossible to unravel phylogenetic relationships within the Coronulida from skeletal morphology alone.

Flying birds flap their wings to generate aerodynamic forces large enough to overcome weight and drag. During this behavior, the forelimbs are displaced and deformed in a complex, coordinated sequence of movements collectively known as the “flight stroke.” Despite an influx of relevant fossil material and new functional insights from extant birds, the historical origin of the avian flight stroke remains poorly resolved. Potential behavioral precursors have been identified primarily on the basis of kinematic resemblance—similarity of movement irrespective of underlying mechanisms. We discuss fundamental issues of motion analysis that are frequently overlooked by paleontologists, and conclude that a purely kinematic approach is insufficient. Consideration of kinetics, the forces responsible for motion, offers a more complete picture of flight stroke evolution. We introduce six kinetic components that interact to determine a limb's trajectory. Phylogenetic mapping reveals that forelimb loading patterns have undergone at least two major transitions on the line from basal archosaur to modern bird. Using this kinematic and kinetic perspective, we offer four specific criteria to help constrain and evaluate competing scenarios for the origin of the avian flight stroke.

Dental microwear analyses of ungulates and other large herbivores rely on correlations of diet and microwear among extant ungulates, primarily ruminants. Microwear is considered a ‘taxon-free’ method of paleodietary analysis. The properties of food are associated with causality of microwear, but the possibility that heritable properties of the consumer (tooth morphologies, masticatory dynamics, enamel mechanical properties, digestive physiologies) may introduce bias is not considered. Using an observer blind method of light microscopy, we examined the distribution of microwear features on the molars of eight species of ruminants and perissodactyls. Grazing and browsing ruminants had statistically different numbers of scratches forming discrete data clusters. Perissodactyls differ in the numbers of scratches and pits but without discrete browser and grazer clusters. Microwear features were distributed homogeneously across ruminant molars and strongly predictive of diet from the labial edge of the molar to the lingual edge. Microwear was heterogeneously distributed across perissodactyl molars with more pits on the labial edge and more scratches on the lingual edge. In perissodactyls, microwear sampled from the labial edge was strongly predictive of diet, while microwear sampled from other areas were not. Discriminant function analyses of microwear assigned individual molars to diets (browser and grazer) and clades (ruminant and perissodactyl) with similar success (70–73%) indicating that phylogeny and diet influence microwear equally. Rhino microwear was more sensitive to clade membership while other perissodactyl microwear was more sensitive to diet. Although it is not clear what heritable variables may phylogenetically bias dental microwear, extant ruminants may not be appropriate models for the microwear of other large herbivores.

Detailed analysis of the stratigraphic ranges of Devonian rugose coral genera within the Old World and Eastern Americas Realms gives new information on faunal extinctions and other bioevents in both realms. Various origination and extinction metrics are calculated from tabulations of occurrences in each stage. The most significant faunal changes were near or at the ends of the Lochkovian and Frasnian stages. The former marks the gradual transition from dominance by Silurian families and genera to the characteristic Devonian coral assemblages; the latter marks the virtual extinction of the Devonian families and genera. Other coral events are related to the two major changes.

The data provide new bases for comparing the histories of the two realms. Most of the events are recorded in both, giving support to previous suggestions that the causes were worldwide. The coral record shows an increase (probably episodic) in environmental deterioration persisting through the Middle Devonian and culminating in extinction at the end of the Frasnian. Eustatic sea level fluctuations may have caused the precursor events and a bolide impact may have caused the end-Frasnian extinction.

A survey of the primary xylem anatomy and dimensions of axes (=“stems”) of Upper Devonian plant petrifaction/compression fossils reveals the following: among genera, the primary xylem strand (1) increasingly deviates from a cylindrical geometry (=haplostele) as both the axis diameter and the nonxylem (“ground”) tissue volume increase and (2) increases in geologically younger specimens. (3) The perimeter length to area ratio of the primary xylem strand (or the primary xylem surface area to volume ratio) is dependent upon axis diameter or the area of the ground tissue. This ratio (4) decreases in evolutionary lineages that involve the appearance of larger plants that maintain a haplostelar anatomy (e.g., the evolution of trimerophytes from a rhyniophyte ancestor) or increases in evolutionary lineages involving the appearance of nonhaplostele anatomy (e.g., the evolution of progymnosperms and pteridophytes from trimerophyte ancestors) and (5) the relative amount of primary xylem to ground tissue increases in stratigraphically younger specimens. These trends are interpreted within the context of (1) correlations between the diameter of axes and the complexity of the primary xylem anatomy reported for some extant pteridophytes and (2) changes associated with the evolution of new primary xylem configurations and organographic relationships. Based on the appearance of new ontogenetic patterns and concomitant changes in the anatomy of the primary xylem strand associated with the evolution of derived plant groups, a stria extrapolation of a size to complexity correlation based on ontogenetic studies to the phyletic level is considered suspect.

There is a well-established discrepancy between paleontological and molecular data regarding the timing of the origin and diversification of placental mammals. Molecular estimates place interordinal diversification dates in the Cretaceous, while no unambiguous crown placental fossils have been found prior to the end-Cretaceous mass extinction. Here, the completeness of the eutherian fossil record through geological time is evaluated to assess the suggestion that a poor fossil record is largely responsible for the difference in estimates of placental origins. The completeness of fossil specimens was measured using the character completeness metric, which quantifies the completeness of fossil taxa as the percentage of phylogenetic characters available to be scored for any given taxon. Our data set comprised 33 published cladistic matrices representing 445 genera, of which 333 were coded at the species level.

There was no significant difference in eutherian completeness across the Cretaceous/Paleogene (K/Pg) boundary. This suggests that the lack of placental mammal fossils in the Cretaceous is not due to a poor fossil record but more likely represents a genuine absence of placental mammals in the Cretaceous. This result supports the “explosive model” of early placental evolution, whereby placental mammals originated around the time of the K/Pg boundary and diversified soon after.

No correlation was found between the completeness pattern observed in this study and those of previous completeness studies on birds and sauropodomorph dinosaurs, suggesting that different factors affect the preservation of these groups. No correlations were found with various isotope proxy measures, but Akaike information criterion analysis found that eutherian character completeness metric scores were best explained by models involving the marine-carbonate strontium-isotope ratios (87Sr/86Sr), suggesting that tectonic activity might play a role in controlling the completeness of the eutherian fossil record.

The marine faunas of tropical America underwent substantial evolutionary turnover in the past 3 to 4 million years in response to changing environmental conditions associated with the rise of the Isthmus of Panama, but the ecological signature of changes within major clades is still poorly understood. Here we analyze the paleoecology of faunal turnover within the family Pectinidae (scallops) over the past 12 Myr. The fossil record for the southwest Caribbean (SWC) is remarkably complete over this interval. Diversity increased from a low of 12 species ca. 10–9 Ma to a maximum of 38 species between 4 and 3 Ma and then declined to 22 species today. In contrast, there are large gaps in the record from the tropical eastern Pacific (TEP) and diversity remained low throughout the past 10 Myr. Both origination and extinction rates in the SWC peaked between 4 and 3 Ma, and remained high until 2–1 Ma, resulting in a 95% species level turnover between 3.5 and 2 Ma. The TEP record was too incomplete for meaningful estimates of origination and extinction rates. All living species within the SWC originated within the last 4 Myr, as evidenced by a sudden jump in Lyellian percentages per faunule from nearly zero up to 100% during this same interval. However, faunules with Lyellian percentages near zero occurred until 1.8 Ma, so that geographic distributions were extraordinarily heterogeneous until final extinction occurred. There were also striking differences in comparative diversity and abundance among major ecological groups of scallops. Free-swimming scallops constituted the most diverse guild throughout most of the last 10 Myr in the SWC, and were always moderately to very abundant. Leptopecten and Argopecten were also highly diverse throughout the late Miocene and early Pliocene, but declined to very few species thereafter. In contrast, byssally attaching scallops gradually increased in both diversity and abundance since their first appearance in our samples from 8–9 Ma and are the most diverse group today. Evolutionary turnover of scallops in the SWC was correlated with strong ecological reorganization of benthic communities that occurred in response to declining productivity and increased development of corals reefs.

Writers as disparate in professional interest as those describing speciation in Drosophila, predicting weather, or supporting creationism have all recently focused on the “sudden” appearance of taxa in the geologic record. In the process, each has cited the punctuated equilibrium argument. Owing to the enormously wide attention which this issue has received since its introduction almost a decade ago, it seems worthwhile to try to place the paleontological and biological evidence in a 1981 perspective. The focus I have chosen is to try to analyze the issue, much as in recent columns in Current Happenings on competitive exclusion, and on larval dispersal and biogeography, in which are put forth alternative explanations for a set of data on a general problem.

I describe a mathematical model of shells that incorporates simple morphometric measurements, which can be obtained easily from sectioned specimens. The model reconstructs form (size and shape) by describing the path in space traversed by, and the changes in dimensions of, a shell aperture during accretionary growth. The model contains parameters that independently affect components of these two aspects of shell form in directions radially away from (or toward) and along a coiling axis. I also describe a computer program, CerioShell, that uses the mathematical model to graphically simulate shell forms. The program permits the viewing of images from different three-dimensional orientations, using various graphical representations, and provides dimensions of the images it generates. Together, the model and the program allow one to investigate the effects that changes in modes of growth would have on shell form.

We use Fourier analysis and related techniques to investigate the question of periodicities in fossil biodiversity. These techniques are able to identify cycles superimposed on the long-term trends of the Phanerozoic. We review prior results and analyze data previously reduced and published. Joint time-series analysis of various reductions of the Sepkoski Data, Paleobiology Database, and Fossil Record 2 indicate the same periodicity in biodiversity of marine animals at 62 Myr. We have not found this periodicity in the terrestrial fossil record. We have found that the signal strength decreases with time because of the accumulation of apparently “resistant” long-lived genera. The existence of a 62-Myr periodicity despite very different treatment of systematic error, particularly sampling-strength biases, in all three major databases strongly argues for its reality in the fossil record.

A marked retardation of dental ontogeny characterizes the family Elephantidae. As a consequence of this retardation, elephant teeth are subject to the forces of mastication, eruption, and progression while still in a developing and pliant stage. As specimens described here illustrate, the mechanical forces are often sufficient to deform the gross morphology of dentitions. Morphological variation in elephant teeth can be regarded as “fabricational noise”—revealing information about the dynamic context in which the teeth develop. Accordingly, dental variation is less species-specific in elephants than in other mammals. The fossil record may comprise fewer species of elephants than is generally believed, and trends inferred to reflect rapid evolution within this family may in fact reflect phenotypic plasticity.

A general model of species dynamics must incorporate the effects of species number on the processes of speciation and extinction. Previous models make specific assumptions about these effects, but do not consider the effects of dynamics of lower level entities on speciation and extinction rates. A hierarchical model is developed which explicitly describes the effects of energy use by species on speciation and extinction rates. The effects of energy use are represented by parameters that characterize the average effects of energy use by each species in the biota on speciation and extinction rates. The dynamics of the model describe a sigmoidal increase in species number over time, as does the logistic model of species dynamics. However, the mechanisms of those dynamics are assumed to be different in the two models. Empirical analysis of a data set on the diversification of fossil Miocene horses suggests that the logistic and hierarchical models have similar descriptive power. The hierarchical model incorporates insights from recent considerations of the nature of a hierarchical theory of biology. Further progress in developing such a theory will depend on the success with which relationships among levels in the biological hierarchy are able to be defined.

Experiments were conducted to test the hypothesis that biomechanical constraints determine the morphology of regular echinoids. Hard-bottom-dwelling Tripneustes gratilla elatensis were transferred to an artifical sandy habitat to evaluate whether the change in substrate affects their height to diameter ratio (H/D). Within 1–2 months their H/D ratio increased significantly. This change was shown to be reversible to some extent. Surgical damage to the ambulacral system of one ray caused inactivation of tubefeet and atrophy of injured ambulacra. Test shape was also affected: the damaged ray was lower, and the nondamaged ambulacra deflected toward the treated one, producing bilateral symmetry as in recorded cases of teratology. Study of T. g. elatensis tetramers showed that while “perfect” tetramery was apparently associated with genetic aberration, “imperfect” tetramery results from mechanical injury at an early ontogenetic stage. Micromorphological study shows that in the longitudinal sutures, normally under tension, long and slender trabeculae develop, associated with long and well-aligned collagenous sutural fibers, while the latitudinal trabeculae and fibers are short and less organized. A mechanical effect is suggested by the oval cross-section of the fiber-anchoring trabeculae. Further, echinoid plates interact like soap bubbles, whereas the entire test behaves like a balloon, fastened to the substrate by the ambulacral tubefeet. All these observations support earlier hypotheses on the biomechanical control of echinoid test growth. A model is proposed in which the expansion of the inner mass, counteracted by the mechanical activity of the ambulacral tubefeet, mesenterial threads, and lantern muscles, affects sutural growth, thus controlling echinoid morphogenesis.

A morphometric survey among regular echinoids reveals an inverse relationship between ambulacral width and relative ambital height. Although both increase of ambulacral width and lowering of ambitus-line are evolutionary trends, it is suggested that they are a response to a mechanical effect. H/D ratio was not related to ambulacral width on the phylogenetic level. It is therefore suggested that the latter correlation is ontogenetically controlled. Aspects of irregular echonoid evolution, such as bilateral symmetry, flattening, and formation of the ambulacral petaloid, also are explained by this model.

Minimum Sample Richness (MSR) is defined as the smallest number of taxa that must be recorded in a sample to achieve a given level of inter-assemblage classification accuracy. MSR is calculated from known or estimated richness and taxonomic similarity. Here we test MSR for strengths and weaknesses by using 167 published mammalian local faunas from the Paleogene and early Neogene of the Quercy and Limagne area (Massif Central, southwestern France), and then apply MSR to 84 Oligo-Miocene faunas from Riversleigh, northwestern Queensland, Australia. In many cases, MSR is able to detect the assemblages in the data set that are potentially too incomplete to be used in a similarity-based comparative taxonomic analysis. The results show that the use of MSR significantly improves the quality of the clustering of fossil assemblages. We conclude that this method can screen sample assemblages that are not representative of their underlying original living communities. Ultimately, it can be used to identify which assemblages require further sampling before being included in a comparative analysis.

Astogenetic trajectories have constrained evolutionary changes in bryozoans. Rates and timing of astogenetic differentiation have been modified for characters defining the morphology of zooids, subcolonies, and colonies. This paper catalogs 46 examples of bryozoan heterochrony, representing all five skeletonized orders. Heterochrony is inferred to have been a pervasive phenomenon in the evolution of Paleozoic stenolaemates, illustrated by 40 examples, 19 of which produced paedomorphosis and 21 peramorphosis. As a consequence, a restricted range of morphologic states has reappeared repetitively as homeomorphies and evolutionary reversals. Large-scale patterns developed across both geologic time and geographic space reflect variation in heterochronic products irrespective of the developmental processes by which they were achieved. Available evidence indicates that smaller, paedomorphic, and more plastic species inhabited onshore, low-diversity areas. Nonheritable plasticity is inferred to be a correlate of early growth stages and paedomorphosis. Taller, generally peramorphic species with damped plasticity are found in higher diversity, offshore regions. Seven key innovations, which first appeared during the early diversification of bryozoan clades, are peramorphic, and recapitulation was a predominant pattern during their Ordovician radiation. Trends in later phylogeny, on the other hand, have favored paedomorphic derived morphologies, as illustrated by 19 of the 32 examples. Recurrent reverse recapitulation suggests that offshore ancestors frequently gave rise to onshore paedomorphs.

Computer models of the tail of Apatosaurus louisae show it could reach supersonic velocities, producing a noise analogous to the “crack” of a bullwhip. Similarity in tail structure suggests this was feasible for other diplodocids, and possibly for unrelated sauropods like Mamenchisaurus and the dicraeosaurids. Lengthening of caudal vertebrae centra between positions 18 and 25 is consistent with adaptation to the stresses generated by such tail motion, as is coossification of vertebrae via diffuse idiopathic skeletal hyperostosis (DISH), which occurs in the same region in about half the specimens. The noise produced may have been used for defense, communication, intraspecific rivalry, or courtship, in which case supersonic “cracking” may have been a sexually dimorphic feature. Comparisons with the club-bearing tails of the sauropods Shunosaurus lii and Omeisaurus tianfuensis show the diplodocid whiplash tail was not well adapted as a direct-impact weapon, bringing the tail-as-weapon hypothesis into doubt.