The purpose of this paper is to explore the possibility and utility of estimating multivariate selection in fossil assemblages, using naticid gastropods as a case study. We used the presence or the absence of a naticid borehole as an index of survival with respect to drilling attacks, enabling us to estimate the multivariate selection gradient exerted by this predator on the shell length and shell thickness of two bivalve genera, Astarte and Spisula. We hypothesized that naticid selection pressure would favor the survival of large, thick-shelled bivalve prey throughout the Cenozoic.

Differential survival of prey was recorded over geologic time using processed bulk assemblages from the Miocene, Pliocene, and Pleistocene Epochs. Multivariate logistic regressions were performed by time period to determine if length and thickness were important factors affecting survival. The direction and magnitude of selection on length and thickness for the two genera ranged from zero (no selection) to large positive or negative values. Only two selection coefficients were significant after sequential Bonferroni corrections: thin-shelled Astarte survived substantially better than thick-shelled Astarte during the Pleistocene (βavggrad = −1.23) and large Spisula survived slightly better than small Spisula during the Miocene Epoch (βavggrad = 0.05).

This is the first study using fossils to calculate multivariate selection gradients. It suggests that naticids were not necessarily strong agents of selection on two traits previously thought to be important to survival of drilling attacks for two of their common prey species. We also show that multivariate selection gradient estimates differ from traditional predation intensity estimates but are superior for estimating the magnitude and direction of natural selection because they use differential mortality of different prey phenotypes rather than just absolute mortality from predation. This work is especially significant for research that involves estimating the relative importance of predation (naticid or otherwise) as an evolutionary force and will be useful for fossil studies where differential survival can be recorded.

This study examines the morphology and function of hindlimb plumage in Archaeopteryx lithographica. Feathers cover the legs of the Berlin specimen, extending from the cranial surface of the tibia and the caudal margins of both tibia and femur. These feathers exhibit features of flight feathers rather than contour feathers, including vane asymmetry, curved shafts, and a self-stabilizing overlap pattern. Many of these features facilitate lift generation in the wings and tail of birds, suggesting that the hindlimbs acted as airfoils. A new reconstruction of Archaeopteryx is presented, in which the hindlimbs form approximately 12% of total airfoil area. Depending upon their orientation, the hindlimbs could have reduced stall speed by up to 6% and turning radius by up to 12%. Presence of the “four-winged” planform in both Archaeopteryx and basal Dromaeosauridae indicates that their common ancestor used fore- and hindlimbs to generate lift. This finding suggests that arboreal parachuting and gliding preceded the evolution of avian flight.

The independent acquisition of high-crowned cheek teeth (hypsodonty) in several ungulate lineages (e.g., camels, equids, rhinoceroses) in the early to middle Miocene of North America has classically been used as an indication that savanna vegetation spread during this time. Implicit in this interpretation is the untested assumption that hypsodonty was an evolutionary response to feeding in open habitats, either due to a change in food source (from browse to graze) or to increased incorporation of airborne grit in the diet. I examined the adaptive explanation for hypsodonty in equids using criteria pertaining to process and pattern of adaptations set up in the comparative-methods literature. Specifically, I tested whether hypsodonty appeared coincident with or just after the spread of open, grass-dominated habitats in the Great Plains of North America.

Phytolith (plant opal) analysis of 99 phytolith assemblages extracted from sediment samples from Montana/Idaho, Nebraska/Wyoming, and Colorado were used to establish the first continuous record of middle Eocene-late Miocene vegetation change in the northern to Central Great Plains. This record was compared with the fossil record of equids from the same area in a phylogenetic framework.

The study showed that habitats dominated by C3 grasses were established in the Central Great Plains by early late Arikareean (≥21.9 Ma), at least 4 Myr prior to the emergence of hypsodont equids (Equinae). Nevertheless, the adaptive hypothesis for hypsodonty in equids could not be rejected, because the earliest savanna-woodlands roughly co-occurred with members of the grade constituting the closest outgroups to Equinae (“Parahippus”) showing mesodont dentition. Explanations for the slow evolution of full hypsodonty may include weak and changing selection pressures and/or phylogenetic inertia. These results suggest that care should be taken when using functional morphology alone to reconstruct habitat change.

The discrete cosine transform (DCT) is a Fourier-related transform widely used in signal processing and well suited to analyzing open outlines such as ammonite ribs. The method is applied here to depict and decipher the ribbing morphospace of a large group of Lower Jurassic ammonites composed of the Oxynoticeratidae and their close ancestors. Because they are clearly associated with buoyancy and/or swimming ability, the usually clearly involute, comparatively smooth and compressed shells of these ammonites may well be misleading taxonomic markers. In this context, quantitative analysis of the ribbing pattern using the DCT may significantly improve our perception of the ornamental patterns expressed within the group. A set of 251 specimens illustrating the worldwide fauna and selected from more than 80 publications is analyzed. Big differences are found in the evolutionary patterns of the two main lineages of Oxynoticeratidae currently accepted in the literature. A previously unsuspected Mediterranean group comprising principally the genus Parasteroceras is identified from its distinctive ornamentation. The northwest European and Mediterranean genera Eparietites, Oxynoticeras, and Parasteroceras do not feature among the American (East Pacific) faunas. This finding calls into question some generally accepted correlations between European and American stratigraphic frameworks. The study shows that the DCT is a valuable tool for discriminating between species within the huge and often puzzling range of ornamental variation of the main genera (e.g., Gleviceras and Radstockiceras).

Evaluating the relative importance of biotic versus abiotic factors in governing macroevolutionary patterns is a central question of paleobiology. Here, we analyzed patterns of changes in global relative abundances and diversity of ecological groups to infer the role of biological interactions as driving evolutionary forces in mid-Mesozoic macrobenthic marine ecosystems. Specifically, we tested the hypothesis of escalation, which states that macroevolutionary patterns were controlled by an increasing pressure exerted by enemies on their victims. Associated with evidence of increasing levels of predation and biogenic sediment reworking (bulldozing) is an increasing representation of predation- and disturbance-resistant groups in the fossil record. In particular, we observe increasing proportions of mobile organisms; a decline of vulnerable epifauna living freely on the substrate; and a trend toward infaunalization of the benthos. These trends were most pronounced in the paleotropics, i.e., the region where biological activity is thought to have been highest. The observation that these changes affected several biotic traits and occurred within independent clades argues against the overriding role of a single key adaptive innovation in causing shifts in ecological abundance. Also, changes in the abiotic environment cannot explain these faunal patterns because of lacking cross-correlations with physico-chemical parameters such as global sea level, climate, and seawater chemistry. We conclude that in marine benthic ecosystems of the mid Mesozoic, enemy-driven evolution, or escalation, was a plausible and important factor.

Spinal ossified tendons are a defining character for Ornithischia, one of the two major clades of dinosaurs. The function of these bony rods has remained a mystery since their first detailed description in 1886. Qualitative approaches to understand ossified tendon function have resulted in different ecological and behavioral interpretations for ornithopod dinosaurs. To evaluate ossified tendon function, this study constructed finite element models of the vertebral column for two ornithopod taxa: Tenontosaurus, which shows the plesiomorphic condition of longitudinally arrayed tendons along the spinous processes, and Brachylophosaurus, which exhibits a lattice of tendons along the spinous processes. Both models predict that ossified tendons stiffened the vertebral column, especially the tail, but the derived lattice of ossified tendons in iguanodontoidean dinosaurs, like Brachylophosaurus, increased spinal stiffness more than the plesiomorphic condition. Caudofemoral muscles that retracted the hindlimb during locomotion attached the femur to the tail in ornithopods. Increased tail stiffness caused by intratendinous ossification may have influenced locomotion by rigidly anchoring M. caudofemoralis longus to the tail, thereby allowing a more forceful retraction of the hindlimb by reducing ventral flexion of the tail during muscle contraction. Ossified tendons may also have been important for storing elastic energy throughout the gait cycle. Moreover, the lattice of ossified tendons stiffened the trunk and tail nearly equally in Brachylophosaurus, indicating the evolution of a postural function by passively supporting the epaxial musculature in maintaining a horizontal vertebral column.

A neurological method for assessment of nasal cavity homologies in extant archosaurs is extended to lambeosaurine hadrosaurids (Dinosauria: Ornithischia) to test functional hypotheses associated with their hypertrophied nasal passages and highly derived cranial crests. The olfactory system and associated cranial nerve pathways that have consistent relationships to soft tissue divisions of the nasal cavity are reconstructed in lambeosaurines on the basis of new paleoneurological data and a comparative phylogenetic approach. The new model of the lambeosaurine olfactory system and nasal cavity shows that a significant portion of nasal cavity proper was located outside the crest cavities and that the primary olfactory region was located rostromedial to the orbits.

All available data indicate that the evolutionary hypertrophy of the nasal cavity occurred predominantly within the non-olfactory nasal vestibule, and that crest development was not causally associated with olfaction. The high level of interspecific and ontogenetic variation in crest shape and nasal vestibule development in lambeosaurine dinosaurs is most consistent with proposed behavioral functions, notably acoustic resonance for intraspecific communication. Despite significant modification to the nasal cavity within Archosauria and its extreme hypertrophy and supraorbital development in Lambeosaurinae, the neural olfactory system and the olfactory region of the nasal cavity proper retain their plesiomorphic positions and associations, suggesting that this system is highly conserved in vertebrate evolution.

Cutan, a resistant non-hydrolyzable aliphatic biopolymer, was first reported in the cuticle of Agave americana and has generally been considered ubiquitous in leaf cuticles along with the structural biopolyester cutin. Because leaves and cuticles in the fossil record almost always have an aliphatic composition, it was argued that selective preservation of cutan played an important role in leaf preservation. However, the analysis of leaves using chemical degradation techniques involving hydrolysis to test for the presence of cutan reveals that it is absent in 16 of 19 taxa (angiosperm and gymnosperm), including many previously reported to contain cutan on the basis of pyrolysis data. Cutan is clearly much less widespread in leaves than previously thought, and its presence or absence does not exert any major bias on the preservation of leaves in the fossil record. In the absence of cutan, other constituents—cutin, plant waxes, and internal plant lipids—are incorporated into the geomacromolecule and contribute to the formation of a resistant aliphatic polymer by in situ polymerization during diagenesis.

Understanding the extent to which the reported fossil record reflects biological history, rather than preservational artifacts or other biasing factors, remains one of the central issues in the interpretation of the history of life on Earth. The development of large interactive paleontological databases, such as the Paleobiology Database (PBDB), allows detailed analyses of the patterns of occurrence, both regionally and globally, of taxa in the fossil record and makes possible testing hypotheses of the controls of the patterns. An analysis of data from the PBDB shows that most genera in the fossil record are rare, whereas a relatively small percentage of taxa account for a disproportionate share of the total occurrences. These ubiquitous taxa tend to be speciose and have long stratigraphic ranges. These patterns of occurrence might represent a true biological signal; it is also possible that they reflect taphonomic processes or are the result of taxonomic practice. In particular, common taxa may be taxonomic wastebaskets, i.e., residual and polyphyletic groups resulting from inadequate systematic attention and/or from taphonomic biases resulting in inadequate specimens being preferentially placed in particular genera. A conceptual model for the development of taxonomic wastebaskets suggests that these taxa should be speciose, widely distributed, common, and old (in terms of year of first description), and that they should be the nominate forms for higher taxa. Our analyses suggest that many of the common taxa in the PBDB are consistent with two or more of these expectations and are thus good candidates for being wastebaskets. These taxa are, however, only a small percentage of total genera. A more detailed examination of one group, early gastropods, indicates that possible wastebaskets still are present in a group that has received much recent systematic work. Given that likely wastebasket taxa are a small fraction of all genera, they probably have little effect on overall temporal patterns of generic richness. Their impact on other types of metrics, such as turnover rates or metrics of community diversity or biogeographic similarity, however, might be quite important.

We describe three giant palaeobatrachid fossil tadpoles of the genus Palaeobatrachus (Nieuwkoop-Faber [NF] stages 60–64) from the Miocene of Randecker Maar, Germany. The largest was 150 mm at the beginning of metamorphosis (stage 60), whereas the smallest was 100 mm and approaching the end of metamorphosis (stage 64). In contrast, normal palaeobatrachid tadpoles and their pipid relatives, both extinct and extant, rarely exceed 60 mm in length. We review here both ecological and pathological conditions that are conducive to the development of gigantism in tadpoles. Tadpoles that lack a thyroid gland become exceptionally large and arrest development at early hindlimb stages (NF stages 53–56). However, the advanced metamorphic stages of the giant Palaeobatrachus tadpoles indicate that they were able to metamorphose, and thus were not athyroid. Environmental factors—pond size and permanence, predators, duration of the growing season—may all contribute to tadpole gigantism in certain extant anuran species. We identify suites of ecological features that distinguish extant anurans with large tadpoles from high-latitude and high-altitude permanent lakes in temperate regions (e.g., certain Rana and Telmatobius) from tropical species, such as Pseudis paradoxa, whose tadpoles normally achieve large size in temporary seasonal ponds. The paleoecology of Randecker Maar suggests that Palaeobatrachus tadpoles lived in a permanent semitropical lake, but one with few predators.

For well over 50 years, the telome theory of Walter Zimmermann has been extremely influential in interpreting the evolutionary history of land plant architecture. Using the “telome/mesome” distinction, and the concept of universal “elementary processes” underlying the change in form in all plants, the theory was an ambitious synthesis based on the proposition that evolutionary change might be understood by a simple set of developmental or evolutionary rules. However, a major problem resides in deciding exactly how assertions of change are to span both developmental and evolutionary domains simultaneously, and, we argue, the theory critically fails testability as a scientific theory. Thus, despite continued popularity for the descriptive terms derived from the theory in evolutionary studies of early land plants, time has come to replace it with a more explicit, testable approach. Presented here is an attempt to clarify perhaps the most important issue raised by the telome theory—whether simple changes in basic developmental processes suffice to describe much of early land plant evolution. Considering the morphology of Silurian–Devonian fossil members, it is hypothesized that early land plants possessed a common set of developmental processes centered on primary growth of shoot apical meristems. Among these were (1) the capacity to monitor and act upon internal physiological status here modeled as “apex strength,” (2) a mechanism for allocation of apex strength in a context-dependent way at each point of branching, (3) a rule for context-dependent apex angle for branches, (4) a largely invariant phyllotaxis unrelated to physiological status, and (5) a simple switch for terminating primary growth, based in part on genetics. Implemented as a set of developmental rules within a simple L-system model, these aspects of primary development in plants determine a sizable range of resultant morphologies, some of which are highly reminiscent of the early fossils. Thus, some support is found, perhaps, for Zimmermann's intuition. However, traditional concepts of growth patterns in plants, including the contrast between epidogenesis and apoxogenesis, require updating. In our reformulation, developmental processes, stated as rules of developmental dynamics, together constitute what we term the plant's developmental state. Using a hypothetico-deductive format, one may hypothesize intrinsic (or genetic) developmental processes that play out as realized developmental activity in specific spatial/temporal contexts, as modified by multiple context factors. The resultant plant morphology is highly dependent on multiple and simultaneous pathway ontogenetic trajectories. Within a likely set of developmental rules reasonably inferred from plant development, some of Zimmermann's elementary processes are perhaps recognizable whereas others are not. Progressively “overtopped” morphologies are easily produced by modifying intrinsic branch allocation. However, even so, the other developmental rules have a profound effect on final architectures. Planate architectures and circination vernation, often treated as special cases by plant morphologists, are perhaps better understood in terms of recurrent or iterative developmental relationships. Much analytic work remains before a completely specified system of rules will emerge. A well-articulated relationship between ontogeny and phylogeny remains fundamentally important in assessing evolutionary change. Fossil and living plants make it abundantly clear that current evolutionary concepts involving modification of a single ontogenetic trajectory from ancestor to descendant need to be greatly expanded into consideration of the entire logical geometry of causation in development. A mechanism for testing is also required that need not wait for complete elucidation at the molecular level. The relative simplicity of plant development, combined with an outstanding fossil record of early members, offers unique opportunities along these lines.

Distinguishing the differential roles of hardpart-input rates and burial rates in the formation of shell beds is important in paleobiologic and sedimentologic studies, because high shelliness can reflect either high population density of shell producers or lack of sediment. The modeling in this paper shows that differences in the relative importance of burial rates and hardpart-input rates lead to distinct patterns with respect to the degree of shelliness and taphonomic alteration in shell beds. Our approach substantially complements other models because it allows computation of both shelliness and assemblage-level alteration. To estimate shelliness, we dissected hardpart-input rates into dead-shell production and shell destruction rates. To estimate assemblage-level alteration, we computed an alteration rate that describes how rapidly shells accrue postmortem damage. Under decreasing burial rates but constant hardpart-input rates, a positive correlation between alteration and shelliness is expected (Kidwell's R-sediment model). In contrast, under decreased destruction rates and/or increased dead-shell production rates and constant burial rates (Kidwell's R-hardpart model), a negative correlation between shelliness and alteration is expected. The contrasting predictions thus provide a theoretical basis for distinguishing whether high shell density in shell beds reflects passive shell accumulation due to a lack of sediment dilution or whether it instead reflects high shell input from a life assemblage. This approach should be applicable for any fossil assemblages that vary in shell density and assemblage-level alteration. An example from the Lower Jurassic of Morocco, which has shell-rich samples less altered than shell-poor samples, suggests that the higher shelliness correlates with higher community-level abundance and lower proportion of juveniles of the main shell producer, supporting the driving role of hardpart-input rates in the origin of the shell-rich samples in this case. This is of significance in paleoecologic analyses because variations in shelliness can directly reflect fluctuations in population density of shell producers.

Attempts to use Fourier methods to quantify ammonoid suture shape have failed to yield robust, repeatable results because sutures are complex curves that violate the assumptions of Fourier mathematics. In particular, sampled sutures are artificially truncated such that the first and last sampled points are not equivalent, the folds are non-stationary, and a position along a horizontal reference axis may map to multiple amplitudes along the suture path. Here I introduce an alternative Fourier method—the windowed short-time Fourier transform (STFT)—that accommodates these characteristics of complex curves. For each suture, digitized landmarks were parameterized using a tangent angle function and then smoothed by convolving with an apodization function. Piece-wise Fourier transforms were then calculated and averaged, resulting in a robust, unique quantification of line morphology. STFT coefficients and estimated power spectra describing the relative weights of harmonics were generated for 576 Paleozoic-basal Triassic ammonoid genera, representative of the range of suture morphotypes. While insensitive to major episodes of taxonomic turnover (Frasnian/Famennian, end-Devonian, and Permian/Triassic extinctions), the summed power data support the previously observed trend toward increasing suture complexity through time. Moreover, partitioning the summed power statistic into harmonic ranges allows novel insight into Paleozoic suture evolution. In particular, the data show significant shifts in the dominant morphotypes during periods of rebound and radiation and suggest that basal Triassic ammonoids possessed unique suture morphotypes when compared with those of the Paleozoic.

The analysis of morphological disparity and of morphospace occupation through the macroevolutionary history of clades is now a major research program in paleobiology, and increasingly so in organismal and comparative biology. Most studies have focused on the relationship between taxonomic diversity and morphological disparity, and on ecological or developmental controls. However, the geographic context of diversification has remained understudied. Here we address geography quantitatively. Diversity, disparity, and paleogeographic dispersion are used to describe the evolutionary history of an extinct echinoderm clade, the class Stylophora (cornutes, mitrates), from the Middle Cambrian to the Middle Devonian (about 128 Myr subdivided into 12 stratigraphic intervals). Taxonomic diversity is estimated from a representative sample including 73.3% of described species and 92.4% of described genera. Stylophoran morphology is quantified on the basis of seven morphometric parameters derived from image analysis of homologous skeletal regions. Three separate principal coordinates analyses (PCO) are performed for thecal outlines, plates from the lower thecal surface, and plates from the upper thecal surface, respectively. PCO scores from these three separate analyses are then used as variables for a single, global, meta-PCO. For each time interval, disparity is calculated as the sum of variance in the multidimensional morphospace defined by the meta-PCO axes. For each time interval, a semiquantitative index of paleogeographic dispersion is calculated, reflecting both global (continental) and local (regional) aspects of dispersion.

Morphospace occupation of cornutes and mitrates is partly overlapping, suggesting some morphologic convergences between the two main stylophoran clades, probably correlated to similar modes of life (e.g., symmetrical cornutes and primitive mitrocystitids). Hierarchical clustering allowed the identification of three main morphological sets (subdivided into 11 subsets) within the global stylophoran morphospace. These morphological sets are used to analyze the spatiotemporal variations of disparity. The initial radiation of stylophorans is characterized by a low diversity and a rapid increase in disparity (Middle Cambrian–Tremadocian). The subsequent diversification involved filling and little expansion of morphospace (Arenig–Middle Ordovician). Finally, both stylophoran diversity and disparity decreased relatively steadily from the Late Ordovician to the Middle Devonian, with the exception of a second (lower) peak in the Early Devonian. Such a pattern is comparable to that of other Paleozoic marine invertebrates such as blastozoans and orthid brachiopods. During the Lower to Middle Ordovician, the most dramatic diversification of stylophorans took place with a paleogeographic dispersion essentially limited to the periphery of Gondwana. In the Late Ordovician, stylophorans steadily extended toward lower paleolatitudes, and new environmental conditions, where some of them radiated, and finally survived the end-Ordovician mass extinction (e.g., anomalocystitids). This pattern of paleobiogeographic dispersion is comparable to that of other examples of Paleozoic groups of marine invertebrates, such as bivalve mollusks.

Intrinsic features of organisms, such as morphology or DNA sequences, and the stratigraphic occurrence of fossils provide distinct evidence of the phylogenetic history of life. Because there is only one true history, we expect the historical signals preserved by these data sets to be similar, and several metrics have been developed to measure the fit of phylogenetic hypotheses to the fossil record. However, a variety of biases affect these metrics, and it is unclear whether they can provide more than an estimate of whether one tree fits the fossil record better than another when used on their own. Here we explore two novel applications of stratigraphic fit metrics when they are used with a combination of phylogeny reconstruction methods that do and do not directly include stratigraphic occurrence data (e.g., cladistics and stratocladistics). In particular, we are interested in whether differences in the stratigraphic fit of cladistic and stratocladistic trees can be used to identify cases in which the stratocladistic results are likely to be more accurate, as well as whether such differences can be used to identify potential problems in the underlying data sets.

Using 550 simulated data sets that were analyzed with cladistics and stratocladistics, we found that the absolute difference in fit to stratigraphy between the results of the two methods was strongly correlated with the probability of character state transition and the accuracy of the stratocladistic results relative to the cladistic results. Completeness of the fossil record and number of taxa included in the analysis were more weakly correlated with stratigraphic fit, and the statistical significance of the differences in fit between the two sets of results did not show a meaningful relationship with improvements in accuracy or potential data problems. These results suggest that measuring the difference in stratigraphic fit between cladistic and stratocladistic trees might be useful for qualitatively estimating whether the addition of stratigraphic data benefited a phylogenetic analysis, and for identifying data sets with high average rates of character state change.

Recent phylogenetic studies of Paleozoic gastropods show the classification provided by the Treatise on Invertebrate Paleontology to include numerous highly polyphyletic taxa. Here we test whether this classification reflects limits on the range of possible designs, general architectural constraints, or common functional solutions for Paleozoic gastropods. Our test evaluates whether superfamilial/subordinal level archetypes as defined by the Treatise evolved more frequently than expected among 626 Late Cambrian–Middle Devonian species. Using a previously established phylogeny and five general shell features (spire angle, exhalent current position, base angle, umbilical width, and apertural inclination) we show that there are fewer gastropod morphotypes than expected given the frequency of change in these features. This is true even after accounting for architectural constraints implied by the data. Moreover, the most common morphotypes include significantly more species and evolved significantly more times than expected. These results imply a set of architectural attractors for lower–middle Paleozoic gastropods, consistent with ecomorphological theory that particular morphotypes are best suited for particular lifestyles. Thus, the Treatise classifications likely reflect these ecomorphological patterns within subclades rather than phylogenetic patterns.