We developed a new method to generate topographic maps of tooth crowns from X-ray synchrotron microtomographic data. Maps are drawn after cervix-plane orientation of tooth image stacks, without the need for a geographic information system. Classical topographic maps with contour lines are complemented by slope maps and angularity maps. Cartography allows precise comparisons of cusps morphologies, and quantification of the directions of cusp axis elongation and slope. Application of this method to muroid rodents with cricetine and murine dental patterns reveals clear-cut differences in cusps morphology that are indicative of the direction of the chewing movement, in agreement with wear facet analyses. Rodents with a murine dental pattern were derived from ancestors with a cricetine pattern, and their origin is associated with important changes in cusp morphology and organization. In order to understand such evolutionary change, our investigation is applied to a sample of extant and fossil muroid rodents that are characterized by either a murine dental plan or a cricetine one, or a dental pattern intermediate between those of cricetines and murines.

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.

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.

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.

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.

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.

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.

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.

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.

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.

Within conifers, active abscission of complete penultimate branch systems is not common and has been described mainly from juveniles. Here I present evidence for the abscission of penultimate branch systems within early so-called walchian conifers—trees with a plagiotropic branching pattern. The specimens studied originate from a middle Early Permian gymnosperm-dominated flora within the middle Clear Fork Group of north-central Texas. Complete branch systems of three walchian conifer morphotypes are preserved; all have pronounced swellings and smooth separation faces at their bases. The source plants grew in a streamside habitat under seasonally dry climatic conditions. The evolution of active branch abscission appears to correspond to an increase in the size of conifers, and this combination potentially contributed to the restructuring of conifer-rich late Paleozoic landscapes. Moreover, trees shedding branch systems and producing abundant litter have the potential to affect the fire regime, which is a factor of evolutionary importance because wildfires must have been a source of frequent biotic disturbance throughout the hyperoxic Early Permian.

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.

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.

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 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.

Physical models of graptolites have been constructed for a range of morphologies, with emphasis on planar, multiramous forms. The models are life size and have the density of a living graptolite, based on the now-established collagenous nature of the periderm and unavoidable assumptions about the amount of extrathecal tissue present in the living colony. These models have been used to test the two main hypotheses of graptolite life habit developed by Bulman, Rickards, Kirk, and others. Testing of graptoloid models in water suggests that many rhabdosome shapes were designed for passive rotation within the water column. This is caused in the models by a variety of modifications, including changes in thecal and stipe orientation, alterations of colony shape and the addition of vanes and hooks. Rotation would only have been useful when the rhabdosome was in directional motion and the frequency of such modifications seems anomalous if no such movement occurred. Thus movement by some means is required, either passively, by changes in buoyancy, or by automobility. Spiralling action would increase the harvesting path of an individual living on a planar, multiramous colony, making this a theoretically advantageous mode of life for the morphology. It would prevent the individual zooids of scandent biserial and uniserial colonies from feeding from the same narrow band of water.

The history of life is punctuated by a number of major transitions in hierarchy, defined here as the degree of nestedness of lower-level individuals within higher-level ones: the combination of single-celled prokaryotic cells to form the first eukaryotic cell, the aggregation of single eukaryotic cells to form complex multicellular organisms, and finally, the association of multicellular organisms to form complex colonial individuals. These transitions together constitute one of the most salient and certain trends in the history of life, in particular, a trend in maximum hierarchical structure, which can be understood as a trend in complexity. This trend could be produced by a biased mechanism, in which increases in hierarchy are more likely than decreases, or by an unbiased one, in which increases and decreases are about equally likely. At stake is whether or not natural selection or some other force acts powerfully over the history of life to drive complexity upward.

Too few major transitions are known to permit rigorous statistical discrimination of trend mechanisms based on these transitions alone. However, the mechanism can be investigated by using “minor transitions” in hierarchy, or, in other words, changes in the degree of individuation of the upper level. This study tests the null hypothesis that the probability (or rate) of increase and decrease in individuation are equal in a phylogenetic context. We found published phylogenetic trees for clades spanning minor transitions across the tree of life and identified changes in character states associated with those minor transitions. We then used both parsimony- and maximum-likelihood-based methods to test for asymmetrical rates of character evolution. Most analyses failed to reject equal rates of hierarchical increase and decrease. In fact, a bias toward decreasing complexity was observed for several clades. These results suggest that no strong tendency exists for hierarchical complexity to increase.

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.

Many authors have proposed scenarios for mass extinctions that consist of multiple pulses or stages, but little work has been done on accounting for the Signor-Lipps effect in such extinction scenarios. Here we introduce a method for computing confidence intervals for the time or stratigraphic distance separating two extinction pulses in a pulsed extinction event, taking into account the incompleteness of the fossil record. We base our method on a flexible likelihood ratio test framework that is able to test whether the fossil record is consistent with any extinction scenario, whether simultaneous, pulsed, or otherwise. As an illustration, we apply our method to a data set on marine invertebrates from the Permo-Triassic boundary of Meishan, China. Using this data set, we show that the fossil record of ostracodes and that of brachiopods are each consistent with simultaneous extinction, and that these two extinction pulses are separated by 720,000 to 1.2 million years with 95% confidence. With appropriate data, our method could also be applied in other situations, such as tests of origination patterns, coordinated stasis, and recovery after a mass extinction.