The unique holotype of the Triassic reptile Sharovipteryx (initially Podopteryx) mirabilis has been reexamined and redescribed, correcting the original account. It is a small lizard-like reptile with an elongate head and densely tubular femur and tibia each longer than the (estimated) intergirdle distance. (Part of the pectoral region has been lost and the remnants remain encased in the matrix.) The matrix retains the impression of portions of the integument, including a patagium that reached from the hindlimbs to the base of the tail. Unlike the original account, the patagium did not extend to the pectoral limbs. Experiments with models indicate that Sharovipteryx could have maintained a shallow glide if the femora were held at a shallow angle to the vertebral column and the tibia and feet extended out at right angles to it, thus stretching the integument. The pectoral limbs (with or without a fringing membrane) might have produced a variable canard; alternatively vertical bending of the tail could have applied drag, each making the glide more stable.

Species abundance data are of vital importance in paleontology, but fossil accumulations invariably represent a biased subset of original source communities. Efforts to quantify taphonomic biases are typically prevented by a lack of independent data on the ecological composition of prehistoric faunas. However, analysis of the continental Holocene record can provide a rare opportunity for independent calibration of fossil abundance patterns. We analyzed a comprehensive data set available for the Holocene avifauna of Sweden to investigate the relationship between species abundance in the recent fossil and zooarchaeological records and in prehistoric source communities, and to characterize the importance of different ecological factors in determining terrestrial vertebrate fossil abundances. The number of assemblages in which species occurred was compared with modern-day species abundance, annual residence, body mass, and ecological realm. Modern-day abundance is only one of several significant predictors of fossil abundance; the strongest predictor is body mass, and Holocene species abundance can be interpreted as a measure of species abundance in source communities for a given size class only. Our study represents one of the only direct attempts to quantify species abundance biases between fossil faunas and source communities, and has general applicability for a wide range of terrestrial vertebrate faunas.

Principal components analysis (PCA) of 21 shell parameters (geometry, sculpture, aperture shape, and suture complexity) in 597 L. Devonian to L. Triassic ammonoid genera (spanning ~166 Myr) shows that eight basic morphotypes appeared within ~20 Myr of the first appearance of ammonoids. With one exception, these morphotypes persisted throughout the Paleozoic, occurring in ~75% of the ~5-Myr time bins used in this study. Morphotypes were not exclusive to particular lineages. Their persistence was not just a product of phylogenetic constraints or longevity, and multiple iterations of the same morphotypes occurred at different times and in different groups. Although mass extinction events severely condensed the range of morphologic variation and taxonomic diversity, the effects were short lived and most extinct morphotypes were usually iterated within 5 Myr. The most important effect of mass extinctions on ammonoid evolutionary history seems to have been their role in large scale taxonomic turnovers; they effectively eliminated previously dominant orders at the Frasnian/Famennian (F/F) (Agoniatitida), the Devonian/Mississippian (D/M) (Clymeniida), and the Permian/Triassic (P/T) (Goniatitida and Prolecanitida) extinctions. Survivors varied from two (P/T) to four (D/M) and five genera (F/F). These events generated sharp reductions in morphologic disparity at the D/M (58%) and at the P/T (59%), but there was a net increase at the F/F (38%). There was no obvious survival bias for particular morphotypes, but 64% are interpreted to have been Nautilus-like nektobenthic. The recurrence of particular combinations of morphology and their strong independence of phylogeny are strong arguments for functional constraint. Intervals between mass extinctions seem to have been relatively static in terms of morphotype numbers, in contrast to numbers of genera. Significant decreases in genus diversity (54%) and morphologic disparity (33%) commenced in the mid-Permian (Wordian/Capitanian boundary), well before the final P/T event.

The stratigraphic distribution of graptoloid species within the upper Middle Ordovician strata of New York State represents a complex pattern of origination, extinction, faunal migration, and fluctuating relative abundances. In particular, the observed patterns of species turnover at graptolite biozone boundaries are apparently strongly correlated with lithofacies, sampling strategies, and the depositional effects of relative sea-level change.

Vertical facies changes that occur within third-order depositional sequences and fourth- or fifth-order parasequences are mirrored by changes in the graptoloid faunal composition. Large-scale faunal turnovers at biozone boundaries tend to occur either at sequence boundaries or at flooding surfaces within sequences (e.g., the base of Highstand System Tracts). The base of the Corynoides americanus and Climacograptus (D.) spiniferus Zones coincide with major onlap events, and the Orthograptus ruedemanni Zone fauna appears just below a Lowstand Systems Tract. Within individual biozones, smaller-scale changes such as the fluctuating relative abundances of graptoloid species coincide with higher-order parasequence cyclicity. These distribution patterns resemble recent computer-generated models for the sequence stratigraphic distribution of hypothetical taxa.

By combining good biogeographic control with a detailed sampling program, we are able to see through the patterns attributable to depositional cyclicity and identify the different components of faunal turnover (migration, speciation, and extinction).

The Scleractinia, which are one of the most important builders of modern reefs, have been considered to have first appeared in the Middle Triassic. Recently, Paleozoic scleractiniamorphs have been reported from both the Ordovician and the Permian, suggesting that the scleractinian-like body plan was already established in the Paleozoic. Those Paleozoic scleractiniamorphs are considered either unsuccessful skeletonized offshoots (extinct experiments) or Paleozoic progenitors of the post-Paleozoic Scleractinia. Permian scleractiniamorphs are characterized by “ancestral” features and have no specific morphologies that deny scleractinian affinities. Molecular phylogenetics also indicate that extant scleractinians are monophyletic and originated long before their Triassic appearance. A Paleozoic origin for the Scleractinia is supported by morphological and molecular phylogenetic data. On the other hand, there is no positive evidence to show that different groups of scleractinians had separate soft-bodied precursors.

The Paleozoic scleractinians evolved within the framework of their basic body plan, and a direct derivation of the Scleractinia from the Rugosa is not probable. The Anthozoa are characterized by a bilaterally symmetrical body plan, which is traditionally considered to have been derived from other radially symmetrical Cnidaria. The problem of the origin of scleractinian body plan may provide a key for deciphering the early anthozoan radiation within the Bilateria. Other examples of Paleozoic Scleractinia and scleractiniamorphs will be found, probably in shallow-water reefal facies or deeper-water communities, bridging the stratigraphic gaps in occurrence and elucidating the origin of the Scleractinia and their body plan.

Our knowledge of the diversity, ecology, and phylogeny of Mesozoic birds has increased significantly during recent decades, yet our understanding of their flight competence remains poor. Wing loading (WL) and aspect ratio (AR) are two aerodynamically relevant parameters, as they relate to energy costs of aerial locomotion and flight maneuverability. They can be calculated in living birds (i.e., Neornithes) from body mass (BM), wingspan (B), and lift surface (S L). However, the estimates for extinct birds can be subject to biases from statistical issues, phylogeny, locomotor adaptations, and diagenetic compaction. Here we develop a sequential approach for generating reliable multivariate models that allow estimation of measurements necessary to determine WL and AR in the main clades of non-neornithine Mesozoic birds. The strength of our predictions is supported by the use of those variables that show similar scaling patterns in modern and stem taxa (i.e., non-neornithine birds) and the similarity of our predictions with measurements obtained from fossils preserving wing outlines. In addition, although our WL and AR values are based on estimates (BM, B, and S L) that have an associated error, there is no cumulative error in their calculation, and both parameters show low prediction errors. Therefore, we present the first taxonomically broad, error-calibrated estimation of these two important aerodynamic parameters in non-neornithine birds. Such estimates show that the WL and AR of the non-neornithine birds here analyzed fall within the range of variation of modern birds (i.e., Neornithes). Our results indicate that most modern flight modes (e.g., continuous flapping, flap and gliding, flap and bounding, thermal soaring) were possible for the wide range of non-neornithine avian taxa; we found no evidence for the presence of dynamic soaring among these early birds.

In this paper we analyze the ontogenies of four species of pachypleurosaur (Reptilia: Sauropterygia) occurring in Triassic-age deposits in the Monte San Giorgio region, Switzerland. Preservation of multiple complete specimens representing a growth series from each taxon allows the comparison of ontogenetic trajectories through a space composed of nine variables important in the evolution of the clade. Trajectories are characterized using the multivariate generalization of the allometry equation and then compared through calculations of the angles between allometry vectors. Individual coefficients of vectors are compared after calculation of bootstrapped confidence intervals. Pachypleurosaur ontogeny is found to be allometric and generally conserved, although significant differences between taxa exist. Shape-disassociated allometric changes characterize the transition between Serpianosaurus and Neusticosaurus, while allometric changes within Neusticosaurus are less significant. N. edwardsii is inferred to have arisen through hypermorphosis. Interpretation of whole-body heterochrony in multivariate analysis is discussed.

Primary vascular architecture of members of the Paleozoic Aneurophytales (Progymnosper-mopsida) is described. This architecture is somewhat more complex but fundamentally similar (homologous) to that of members of the Trimerophytina, putative ancestors of aneurophytes. It is suggested that the presence of complex stelar morphology in aneurophytes was epiphenomenal, a passive result of changes in growth and development in a trimerophyte-like ancestor. Specifically, I suggest that the evolutionary transformation in primary vascular architecture from haplostele to ribbed protostele was a direct consequence of changes that affected the vertical spacing and degree of organization of lateral appendages in early vascular plants. This view is in sharp contrast to adaptationist explanations of change in stelar morphology expressed by other authors and provides an example of non-adaptive change in evolution.

The floral community along South Africa's southwest coast today is dominated by shrubby strandveld, renosterveld, and coastal fynbos vegetation. The grass family (Poaceae), represented primarily by C3 taxa, is scarce by comparison. Nevertheless, grass has a long history along this coast, as indicated by the presence of ∼5-million-year-old C3 grass pollen and phytoliths in sediments at the fossil locality of Langebaanweg E Quarry. Because the pollen and phytoliths of other plant families, including fynbos, have also been found, it has been difficult to determine whether grass was scarce or abundant in this environment. In order to shed light on this issue, I analyzed the dental mesowear of the E Quarry bovids. Results indicate that only one (Simatherium demissum) of seven analyzed species was a grazer. These compare well with the results of a microwear texture analysis, which indicate that none of the seven analyzed species were obligate grazers. These two studies point strongly toward a heavily wooded environment and not one that was dominated by grass. Although a conventional dental microwear analysis did identify three out of seven E Quarry bovid species as grazers (Bed3aN Damalacra, Kobus subdolus, and S. demissum), only S. demissum probably actually was a grazer. I suggest that the grazer signal exhibited by the other two bovid samples indicate that these species were taking advantage of a spike in grass abundance, probably during the winter growth season.

Evolutionary trends observed over large clades have the potential to mask underlying trends that occur within their constituent subclades. A recent study of encephalization in the Caniformia (Carnivora, Mammalia) found evidence for an abrupt increase in median log-encephalization quotients (logEQs), indicating higher brain volume relative to body mass, at the end-Miocene, but gradual increase in the variance of logEQs. In this study, new endocranial volume estimates for fossil taxa in the well-sampled caniform subclade Canidae are reported. Using the encephalization data for the Canidae, hypotheses of evolution in encephalization allometries were tested with respect to canid phylogeny. The Akaike Information Criterion and likelihood ratios recovered support for a preferred hypothesis of the evolution of canid encephalization, which proposed two distinct allometric relationships: (1) a plesiomorphic grade of encephalization in the subfamilies Hesperocyoninae and Borophaginae and the paraphyletic canine genus Leptocyon, and (2) an apomorphic grade in the crown radiation of Caninae. This defines a shift in to higher encephalization, but without an associated change in the variance around the allometry. Increased canid encephalization coincides with a reorganization of the brain and the observed trend may reflect the evolution of complex social behavior in this clade.

Despite a global fossil record, Metatheria are now largely restricted to Australasia and South America. Most metatherian paleodiversity studies to date are limited to particular subclades, time intervals, and/or regions, and few consider uneven sampling. Here, we present a comprehensive new data set on metatherian fossil occurrences (Barremian to end Pliocene). These data are analyzed using standard rarefaction and shareholder quorum subsampling (including a new protocol for handling Lagerstätte-like localities).

Global metatherian diversity was lowest during the Cretaceous, and increased sharply in the Paleocene, when the South American record begins. Global and South American diversity rose in the early Eocene then fell in the late Eocene, in contrast to the North American pattern. In the Oligocene, diversity declined in the Americas, but this was more than offset by Oligocene radiations in Australia. Diversity continued to decrease in Laurasia, with final representatives in North America (excluding the later entry of Didelphis virginiana) and Europe in the early Miocene, and Asia in the middle Miocene. Global metatherian diversity appears to have peaked in the early Miocene, especially in Australia. Following a trough in the late Miocene, the Pliocene saw another increase in global diversity. By this time, metatherian biogeographic distribution had essentially contracted to that of today.

Comparison of the raw and sampling-corrected diversity estimates, coupled with evaluation of “coverage” and number of prolific sites, demonstrates that the metatherian fossil record is spatially and temporally extremely patchy. Therefore, assessments of macroevolutionary patterns based on the raw fossil record (as in most previous studies) are inadvisable.

Biomechanical models illustrate how the principles of physics and physiology determine function in organisms, allowing ecological inferences and functional predictions to be based on morphology. Dynamic lever and linkage models of the mechanisms of the jaw and skull during feeding in fishes predict function from morphology and have been used to compare the feeding biomechanics of diverse fish groups, including fossil taxa, and to test ideas in ecological morphology. Here we perform detailed computational modeling of the four-bar linkage mechanism in the skull and jaw systems of Dunkleosteus terrelli, using software that accepts landmark morphological data to simulate the movements and mechanics of the skull and jaws during prey capture. The linkage system is based on the quadrate and cranio-thoracic joints: Cranial elevation around the cranio-thoracic joint forces the quadrate joint forward, which, coupled with a jaw depressor muscle connecting the jaw to the thoracic shield, causes the jaw to rotate downward during skull expansion. Results show a high speed transmission for jaw opening, producing a rapid expansion phase similar to that in modern fishes that use suction during prey capture. During jaw closing, the model computes jaw and skull rotation and a series of mechanical metrics including effective mechanical advantage of the jaw lever and kinematic transmission of the skull linkage system. Estimates of muscle cross-sectional area based on the largest of five specimens analyzed allow the bite force and strike speed to be estimated. Jaw-closing muscles of Dunkleosteus powered an extraordinarily strong bite, with an estimated maximal bite force of over 6000 N at the jaw tip and more than 7400 N at the rear dental plates, for a large individual (10 m total length). This bite force capability is among the most powerful bites in animals. The combination of rapid gape expansion and powerful bite meant that Dunkleosteus terrelli could both catch elusive prey and penetrate protective armor, allowing this apex predator to potentially eat anything in its ecosystem, including other placoderms.

Recent excavations of Ediacaran assemblages have revealed striking bed-to-bed variation in diversity–abundance structure, offering potential insight into the ecology and taphonomy of these poorly understood early Metazoan ecosystems. Here we compare faunal variability in Ediacaran assemblages to that of younger benthic assemblages, both fossil and modern. We decompose the diversity of local assemblages into within-collection (α) and among-collection (β) components and show that β diversity in Ediacaran assemblages is unusually high relative to younger assemblages. Average between-bed ecological dissimilarities in the Phanerozoic fossil record are comparable to within-habitat dissimilarities typically observed over meter to kilometer scales in modern benthic marine habitats, but dissimilarities in Ediacaran assemblages are comparable to those typically observed over 10–100 km scales in modern habitats. We suggest that the unusually variable diversity–abundance structure of Ediacaran assemblages is due both to their preservation as near snapshots of benthic communities and to original ecological differences, in particular the paucity of motile taxa and the near lack of predation and infaunalization.

Determining whether a species has gone extinct is a central problem in both paleobiology and conservation biology. Past literature has mostly employed equations that yield confidence intervals around the endpoints of temporal ranges. These frequentist methods calculate the chance of not having seen a species lately given that it is still alive (a conditional probability). However, any reasonable person would instead want to know the chance that a species is extinct given that it has not been seen (the posterior probability). Here, I present a simple Bayesian equation that estimates posteriors. It uninterestingly assumes that the sampling probability equals the frequency of sightings within the range. It interestingly sets the prior probability of going extinct during any one time interval (E) by assuming that extinction is an exponential decay process and there is a 50% chance a species has gone extinct by the end of its observed range. The range is first adjusted for undersampling by using a routine equation. Bayes' theorem is then used to compute the posterior for interval 1 (ε1), which becomes the prior for interval 2. The following posterior ε2 again incorporates E because extinction might have happened instead during interval 2. The posteriors are called “creeping-shadow-of-a-doubt values” to emphasize the uniquely iterative nature of the calculation. Simulations show that the method is highly accurate and precise given moderate to high sampling probabilities and otherwise conservative, robust to random variation in sampling, and able to detect extinction pulses after a short lag. Improving the method by having it consider clustering of sightings makes it highly resilient to trends in sampling. Example calculations involving recently extinct Costa Rican frogs and Maastrichtian ammonites show that the method helps to evaluate the status of critically endangered species and identify species likely to have gone extinct below some stratigraphic horizon.

A computer simulation was developed which estimates the effects of differential preservation on the value of taxonomic diversity in fossil assemblages. Taxonomic diversity was measured with the Shannon-Weaver index, H′ and with evenness, J. Preservation markedly alters quantitative measurements of taxonomic diversity. The degree of alteration can, however, be predicted on the basis of the species-abundance distribution and on the distributions of species of varying preservability. The indices of diversity H′ and J measure different aspects of diversity, but respond to differential preservation in a similar manner. Evenness, J, is recommended for paleoecologic use. The computer model can be used to revise fossil diversity data. A simple form of such a model has been used, and suggests that some previous interpretations of paleoecologic diversity may be in error.

Standard techniques of energy exchange analysis are applied to modelling the thermal regimes of several species of sailback pelycosaurs. Of particular interest is the role played by the sail in thermoregulation. Although the sail did increase the rate at which the model could warm up in the morning, its effectiveness fell short of previous estimates. The sail increased the daytime internal temperature by typically 3°C to 6°C. The more massive subjects had the highest temperature increases owing partly to their higher thermal inertia and partly to their disproportionately large sails. The sail had no impact upon the nocturnal minimums in temperature.

Overheating does not appear to have been a significant problem in the Permian environment, particularly in view of the high thermal inertia of the subjects modeled. When overheating occurred, the sail was of limited value for dumping excess heat.

Amino acid analyses of undecalcified samples of fossil crocodile and rhinoceros enamel and dentin from mature teeth revealed that the total protein content of these mineralized fossil tissues varied from ~0.01–0.007% by weight. Except in one instance, amino acid analyses of the enamel proteins revealed them to be free of collagen and to have an amino acid composition similar to the proteins obtained from the enamel of mature modern vertebrates. Molecular sieving of the acid soluble enamel proteins demonstrated that the components consisted principally of small peptides and free amino acids, as in the enamel of modern vertebrates.

Based on the presence of hydroxyproline (hyp) and hydroxylysine (hyl), collagen was detected in undecalcified mature dentin of fossil rhinoceros, but not in undecalcified crocodile dentin. It was only by sequential extraction procedures that the presence of collagen in the dentin of fossil crocodile was established, emphasizing the utility and importance of analyzing the soluble components in specific extracts of the fossil. Based on these data and the concentrations of hyp and hyl in the material solubilized by the various solvents, the dentin of fossil rhinoceros contained considerably more collagen per weight and as a percentage of the total protein in the dentin than did the dentin of fossil crocodile.

As with modern dentin, EDTA and dilute acid solubilize the noncollagenous proteins and peptides found in fossil enamel and dentin, some of which contain O-phosphoserine [Ser(P)], an amino acid unique to mineralized connective tissues. Similar to recent reported findings from fossil bone, less of the original content of the noncollagenous proteins, including those phosphorylated, is degraded and removed from the enamel and dentin during fossilization than the percentage of dentinal collagen and the nonphosphorylated domains of the enamel proteins which are removed. This selective resistance to degradation and removal of the noncollagenous proteins, including phosphoproteins, may reflect the strong interaction of these proteins with the mineral phase of fossilized tissues. The amount and close packing of the inorganic crystals may also inhibit the interaction of the proteins in the interior of the enamel and dentin with the geochemical environment.

Cases of convergent evolution, particularly within ecomorphological contexts, are instructive in identifying universally adaptive morphological features across clades. Tracing of evolutionary pathways by which ecomorphological convergence takes place can further reveal mechanisms of adaptation, which may be strongly influenced by phylogeny. Ecomorphologies of carnivorous mammals represent some of the most outstanding cases of convergent evolution in the Cenozoic radiation of mammals. This study examined patterns of cranial shape change in the dog (Canidae) and hyena (Hyaenidae) families, in order to compare the evolutionary pathways that led to the independent specialization of bone-cracking hypercarnivores within each clade. Geometric morphometrics analyses of cranial shape in fossil hyaenids and borophagine canids provided evidence for deep-time convergence in morphological pathways toward the independent evolution of derived bone-crackers. Both clades contained stem members with plesiomorphic generalist/omnivore cranial shapes, which evolved into doglike species along parallel pathways of shape change. The evolution of specialized bone-crackers from these doglike forms, however, continued under the constraint of a full cheek dentition and restriction on rostrum length reduction in canids, but not hyaenids. Functionally, phylogenetic constraint may have limited borophagine canids to crack bones principally with their carnassial instead of the third premolar as in hyaenids, but other cranial shape changes associated with durophagy nevertheless evolved in parallel in the two lineages. Size allometry was not a major factor in cranial shape evolution in either lineage, supporting the interpretation of functional demands as drivers for the observed convergence. The comparison between borophagines and hyaenids showed that differential effects of alternative functional “solutions” that arise during morphological evolution may be multiplied with processes of the “macroevolutionary ratchet” already in place to further limit the evolutionary pathways available to specialized lineages.

Fossil species of the family Hyaenidae represent a wide range of ecomorphological diversity not observed in living representatives of this carnivoran group. Among them, the cursorial meat-and-bone specialists are of particular interest not only because they were the most cursorial of the hyaenids, but also because they were the only members of this family to spread into the New World. Here we conduct a functional morphological analysis of the cranium of the cursorial meat-and-bone specialist Chasmaporthetes lunensis by using finite element modeling to compare it with the living Crocuta crocuta, a well-known bone-cracking carnivoran. As found with previous finite element studies on hyaenid crania, the cranium of C. lunensis is not differentially adapted for stress dissipation between the bone-cracking and meat-shearing teeth. A smaller occlusal surface on the more slender P3 cusp of C. lunensis allowed this species to use less bite force to crack a comparably-sized bone relative to C. crocuta, but higher muscle masses in the latter probably allow it to process larger food items. We use two indices, the stress slope and the bone-cracking index, to show that C. lunensis has a well-adapted cranium for stress dissipation given its size, but the main stresses placed on its cranium might have been more from subduing prey and less from cracking bones. Throughout the Cenozoic, other carnivores besides hyaenids convergently evolved similar morphologies, including domed frontal regions, suggesting an adaptive value for a repetitive mosaic of features. Our analyses add support to the hypothesis that bone-cracking adaptations are a complex model that has evolved convergently several times across different carnivoran families, and these predictable morphologies may evolve along a common gradient of functionality that is likely to be under strong adaptive control.

Edge-drilling is an unusual predation pattern in which a predatory gastropod drills a hole on the commissure between the valves of a bivalve. Although it is faster than wall drilling, it involves the potential risk of amputating the drilling organ. We therefore hypothesize that this risky strategy is advantageous only in environments where predators face high competition or predation pressure while feeding. The high frequency of edge-drilling (EDF, relative to the total number of drilled valves) in a diverse Recent bivalve assemblage from the Red Sea enables us to test this hypothesis, predicting (1) a low EDF in infaunal groups, (2) a high EDF in bivalves with elongated shape, (3) high incidence of edge-drilling in groups showing a high wall-drilling frequency, and (4) high EDF in shallow habitats. We evaluate these predictions based on >15,000 bivalve specimens. Among ecological attributes, we found substrate affinity and predation intensity of a species to be good predictors of edge-drilling incidence. Infaunal taxa with high length/width ratio have a low EDF, in accordance with our predictions. Predation intensity is also a significant predictor of edge-drilling; groups with high predation intensity show higher incidence of edge-drilling, confirming our prediction. Although water depth fails to show any significant effect on EDF, this analysis generally supports the high-risk hypothesis of edge-drilling incidence because shallow depths have considerable microhabitat variability. Classically the drill hole site selection has often been linked to predatory behavior. Our study indicates that prey attributes are also crucial in dictating the behavioral traits of a driller such as site selection. This calls for considering such details of the prey to fully understand predation in modern and fossil habitats. Moreover, this perspective is important for tackling the longstanding riddle of the limited temporal and spatial distribution of edge-drilling.