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

Discovery of several new specimens of the gigantic Eocene ground bird Diatryma gigantea from the Willwood Formation of northwestern Wyoming, has prompted an analysis of its feeding apparatus and an assessment of the mode of life of this unusual bird. Diatryma exhibits many of the features predicted by biomechanical models to occur in animals delivering large dorsoventral bite forces. Similarly, the mandible of Diatryma, which was modeled as a curved beam, appears well equipped to withstand such forces, especially if they were applied asymmetrically. Interpretation of these size-independent biomechanical properties in light of the large absolute skull size of Diatryma suggests a formidable feeding apparatus. The absence of modern analogues complicates the determination of just how this unique skull morphology correlates with diet. Suggestions that Diatryma was an herbivore seem improbable in that they require the postulation of excessively high safety factors in the construction of the skull. The traditional hypothesis of Diatryma as a carnivorous bird accords as well or better with the data at hand. Carnivory raises the probability of “accidental” encounter with bones, thus explaining the high safety factors. In fact, the skull and mandible of Diatryma are so massive that bone crushing may have been an important behavior. Diatryma could have been a scavenger. However, limb allometry and phylogenetic interpretation of limb proportions call into question the picture of Diatryma as a slow, plodding graviportal animal, suggesting that active predation was within its behavioral repertoire.

Under favorable circumstances, biogeographic and biostratigraphic data can be combined to identify accurately the time and place of origin of a given taxon, and to reconstruct the pattern of its subsequent radiation. This study considers the dendrasterid sand dollars, which are abundant today along the Pacific Coast of North America. The Neogene sand dollar record in this region is particularly good; in fact, sand dollars have traditionally been used as provincial index fossils.

The dendrasterids originated in central California at the end of the Miocene; the oldest forms are dated at about 6.0–6.5 Ma. They spread south to Baja California during the Pliocene, and then north to Alaska during the Quaternary. This historical pattern is not an artifact of the record; it is consistent with independent paleogeographic evidence. The dendrasterids supplanted an older Mio-Pliocene sand dollar fauna; they are now completely dominant in the temperate coastal waters of the northeastern Pacific. They have reached this position in less than 7 m.y. since their first local appearance. The rapid rise of dendrasterids could be related to their aberrant morphology and behavior; these adaptations allow dendrasterids to suspension-feed, in a manner unique among living echinoids.

Dendrasterids are characterized by “eccentric” test morphologies. Even the oldest species are highly eccentric; transitional forms are unknown. The first dendrasterids appear suddenly in the provincial “Jacalitos Stage,” above an unconformity which represents no more than about 1 m.y. They do not occur in the underlying units, although other fossil sand dollars are abundant. The dendrasterids may have arisen rapidly, through a heterochronic change in the development of older, noneccentric forms. Recent ontogenetic studies have documented the feasibility of this process.

Using the diameters of the channels at branch points, we quantitatively test three alternative models of the function of astrorhizae in stromatoporoids. The distribution of diameters at branch points is significantly different from the distribution that would be predicted from models of either a diffusive function or a bulk-flow system in which resistance to flow was constant at all levels of the branching hierarchy. The distribution of channel diameters is virtually identical to that predicted by a model (Murray's law) that simultaneously minimizes resistance to flow and some volume-related cost function. Astrorhizae thus carried a bulk flow of fluid and can be inferred to have been lined with cellular elements; the exchange sites associated with the fluid-transport system were distributed throughout the soft tissues of the stromatoporoid animal. The most parsimonious hypothesis of function, that the fluid-transport system was associated with suspension feeding, implies strong similarities between the structure of the stromatoporoid animal and living sponges.

Many organisms continue to grow their skeletons throughout ontogeny. In the shells of molluscs, protists, and brachiopods and in bovid horns, accretionary spiral growth provides a detailed and continuous growth history. Although a shell may be described as a single static form, the overall morphology is a summation of the ongoing accretionary process. For this reason, an explicitly ontogenetic characterization of form provides insight into the final form achieved. Analysis of landmark transformations offers direct access to major components of morphological variation, both among adult individuals and through an individual's ontogeny. Parameters of preconceived, abstract geometric models can also be used to characterize morphological variation, but there is no guarantee that these parameters will coincide with the major features of shape variation.

In order to locate landmarks at equivalent ontogenetic stages, features that indicate ontogenetic stage of coiled forms must be identified (e.g., growth increments, age, size, whorls). The gastropod Epitonium (Nitidiscala) tinctum exhibits prominent varices that provide landmark locations throughout ontogeny. Recent specimens of this species were obtained from three localities in Baja, Mexico. The morphological variation among individuals, treated as whole shells and within individual ontogenies, was analyzed using shape coordinates of landmark configurations. Deformation of shape is expressed in the uniform and nonuniform shape subspaces. The empirical components of shape variation found are similar to those generated by two parameters of an equiangular spiral: θ, the angle between consecutive varices, and W, the whorl expansion rate. The distribution of individuals is examined within morphospaces constructed from these shape features.

Three scales of analysis are necessary to characterize adequately the shape variation within and among specimens. The smallest scale is equivalent to increment-by-increment changes in θ and W. The middle scale comprises variation equivalent to whorls resulting from systematic changes in θ and W during an individual's ontogeny. Finally, there is the overall ontogenetic trajectory. Mean shape must be a function of initial shape and ontogenetic trajectory in shape. Mean forms that are found to have similar shapes at the same arbitrary growth increment may achieve that shape in different ways.

Convergent evolution of hypercarnivorous adaptations in canids has occurred a number of times in the last 40 m.y. among distantly related taxa. The adaptations include an increase in carnassial blade length, reduction or loss of post-carnassial molars, and transformation of the talonid of the lower first molar from a basinlike depression into a trenchant, bladelike cusp. Although the diversity of these specialized canids is typically low in past and present communities, it was unusually high during the Late Oligocene of North America and the Pleistocene of South America. These two comparable events provide an opportunity for exploring possible causes of the evolution of hypercarnivory in canids. Plots of generic diversity against time for North American predators reveal a roughly inverse relationship between the number of hypercarnivorous canid taxa and the numbers of other hypercarnivores, such as creodonts, nimravids, mustelids, and amphicyonids. Similarly, the radiation of hypercarnivorous canids in South America occurred at a time of relatively low diversity of other hypercarnivores. Analysis of trophic diversity within the North American carnivore paleoguild before, during, and after the Late Oligocene reveals considerable taxonomic turnover among carnivores because of immigration and speciation. Late Oligocene hypercarnivorous canids appear to have been replaced first by amphicyonids and large mustelids, and then by felids.

Despite the repeated tendency of canids to evolve adaptations for hypercarnivory, a canid has yet to appear that is completely catlike, that is, without any post-carnassial molars. This possible constraint on morphological evolution in canids is argued to have resulted, paradoxically, in increased flexibility over evolutionary time and a great potential for rapid diversification and clade survivorship. Finally, it is suggested that the iterative pattern of specialization of the lower molars for meat-slicing that is seen in all families of carnivores, past and present, is probably a result of intraspecific competition for food, perhaps among littermates. This intraspecific selective force is countered by competition among species, since there are limits on the number of sympatric hypercarnivorous species within a single community.

Evolutionary progress is a trend that relaxes trade-off rules. It begins with the evolution of a key adaptation. It continues with the spread of the key adaptation as the clade that contains it replaces some older clade that lacks it. Key adaptations are those that allow for improvement in at least one organismal function at a reduced fitness cost in other functions.

Replacement almost certainly involves more than pure chance. It may not often involve competitive extinction. Instead, species from the new clade produce new species to replace already extinct species from the old clade. The key adaptation gives them a higher competitive speciation rate than old-clade sources of replacement. The process, termed incumbent replacement, proceeds at a rate limited by extinction rate. Thus, replacement often seems linked to mass extinction events.

The incumbent-replacement hypothesis explains what we know about the replacement of straight-neck turtles (Amphichelydia) by those that can flex their necks and protect their heads in their shells. This replacement occurred four or five times in different biotic provinces. It happened as long ago as the Cretaceous in Eurasia, and as recently as the Pleistocene in mainland Australia. It was accomplished in Gondwanaland by turtles flexing their necks sideways (Pleurodira), and in the north by those flexing their necks into an S-curve (Cryptodira). As is typical of replacements, amphichelydian replacement took millions of years to accomplish wherever it occurred, and much of it in North America took place in a burst associated with and immediately subsequent to a mass extinction.

Studies of the fates of clades during mass extinctions at the end of the Cretaceous and near the Cambrian-Ordovician boundary have indicated that geographically widespread groups have a much higher probability of survival than narrowly distributed groups. If biogeographic distribution does play a significant role in influencing the outcome of extinction events, then geographic variability in extinction intensity should be mirrored by patterns of endemism. A comparison of data for the terminal Cambrian extinction indicates that survival of trilobite families in Kazakhstan was significantly greater than in North America. As predicted, the Kazakh sequence was characterized by a significantly larger number of pandemic families. The Baltic Province, which includes Scandinavia, England, and Wales, was composed entirely of pandemic groups and did not suffer any trilobite extinction at the family level. The results are consistent with the suggestion that latitudinal variation in extinction magnitude may be the result of differences in biogeographic structure.

A kill curve for Phanerozoic species is developed from an analysis of the stratigraphic ranges of 17,621 genera, as compiled by Sepkoski. The kill curve shows that a typical species' risk of extinction varies greatly, with most time intervals being characterized by very low risk. The mean extinction rate of 0.25/m.y. is thus a mixture of long periods of negligible extinction and occasional pulses of much higher rate. Because the kill curve is merely a description of the fossil record, it does not speak directly to the causes of extinction. The kill curve may be useful, however, to limit choices of extinction mechanisms.

The impact of the variations in the chemical composition of higher vascular plant cuticles on their fossil record is usually not considered in paleobotanical and, more particularly, taphonomic studies. Here we address the subject with reference to the chemical characterization of insoluble cuticular matrices of a large variety of recent and fossil cuticles. The cuticles were analyzed using Curie-point pyrolysis-gas chromatographic techniques. Cuticular matrices of extant higher plants consist either of the biopolyester cutin, the insoluble, non-hydrolyzable polymethylenic biopolymer cutan, or a mixture of both biopolymers. In fossil cuticles an additional cuticular matrix type consisting of cutan and cutin-derived material is recognized. On the basis of the variations in their chemical composition and the different behavior of the cuticular constituents (viz., cutin and cutan) during diagenesis, it is concluded that the paleobotanical record of cuticles will be biased toward taxa originally having a significant amount of cutan in their cuticular matrix.

The stratigraphic framework for Triassic and Early Jurassic continental strata has greatly changed in recent years. These revised correlations necessitate a review of traditional views of early Mesozoic continental faunal succession and biogeography. We have examined the relationship between tetrapod distribution and paleogeographic context during the Triassic and Early Jurassic on the basis of a data base comprising updated faunal lists for major early Mesozoic assemblages of continental tetrapods. Analysis of these data supports the hypothesis that there were few barriers to biotic interchange among continental tetrapods throughout the Triassic and Early Jurassic. Early Mesozoic tetrapod assemblages are dominated by widely distributed, often cosmopolitan families. Late Triassic patterns of latitudinal variation among tetrapod assemblages appear to be correlated to those seen among terrestrial plants and contrast with the extremely uniform distribution of Early Jurassic continental biotas.

Colony growth form in some Silurian heliolitid corals is analyzed by the measurement of their shape in profile. Data are presented for seven species, Stelliporella parvistella, Heliolites interstinctus, H. megastoma, H. daintreei, H. spongodes, Propora tubulata, and Plasmopora scita from three localities in Gotland, Sweden, and three localities in England. Intraspecific growth-form variation is presented on triangle diagrams. These plots allow variation to be compared between species present at each locality and between localities for each species. Results indicate that the overall potential for growth-form variation is genetically controlled and that levels of response to environmental stimuli may differ markedly between species. Stelliporella parvistella is a very plastic species, the only one developing branching growth in addition to other growth forms. Heliolites interstinctus is much less variable, dominantly tabular, domal, and low bulbous in form, but demonstrates a similar response. Propora tubulata has a tightly constrained bulbous growth form that shows little variation between localities. The other species are represented by few specimens, most of which parallel H. interstinctus. The likely moderating influences of light levels, substrate type, sedimentation rate, energy levels, and other variables on growth-form variation and species range are considered. The main environmental factor including ecophenotypic response is concluded to be sedimentation rate. A close correlation between this factor and growth form in S. parvistella indicates that form in this species is a particularly sensitive indicator of sedimentation rate and substrate conditions. No simple equations can be made between specific environments and one particular growth form in these corals.

Taxon-age distributions were compiled for families of marine animals surviving or becoming extinct in each stage of the Phanerozoic. I demonstrate, through the use of a modified bootstrap analysis, that there is no difference between the longevity of families becoming extinct during times of background extinction and times of mass extinction. In both mass and background extinction intervals the mean age of families that become extinct is 2 standard deviations below the geometric mean taxon age of families available for extinction. Young families are more susceptible to extinction, perhaps as the result of lower species richness or of occupying a smaller geographic range. There is no tendency during mass extinctions toward loss of families with different taxon ages other than those that become extinct during background times. Thus, in terms of family survival, mass extinction appears to be an exaggeration of processes of background extinction.

Problems of stratigraphic completeness and poor temporal resolution make analysis of faunal change in terrestrial sequences difficult. The fluvial Neogene Siwalik formations of India and Pakistan are an exception. They contain a long vertebrate record and have good chronostratigraphic control, making it possible to assess the influence of biotic interchange on Siwalik fossil communities. In Pakistan, the interval between 18 and 7 Ma has been most intensively studied and changes in diversity and relative abundance of ruminant artiodactyls and muroid rodents are documented with temporal resolution of 200,000 years. Within this interval, diversity varies considerably, including an abrupt rise in species number between 15 and 13 Ma, followed by a decline in ruminant diversity after 12 Ma and a decline in muroid diversity in two steps at 13 and 10 Ma. Significant changes in relative abundance of taxa include an increase in bovids between 16.5 and 15 Ma, a decrease in tragulids after 9 Ma, and a very abrupt increase in murids at 12 Ma. Megacricetodontine rodents also decrease significantly at 12 Ma, and smaller declines are recorded among myocricetodontine and copemyine rodents after 16 Ma. An increase of dendromurine rodents at 15.5 Ma is also observed. There is also a trend of progressive size increase among giraffoids and bovids throughout the sequence.

We have also investigated relationships between biotic interchange and diversity, body size, and relative abundance, concluding that (1) the rapid increase in ruminant and muroid diversity was largely due to immigration, whereas in situ speciation had only a secondary role; (2) during intervals of increasing diversity, resident lineages did not have higher than average rates of in situ speciation; (3) during intervals with rising diversity, greater extinction did not accompany increased immigration; (4) during intervals with falling diversity, there may have been greater extinction in recently invading lineages; and (5) change in diversity was independent of changes in relative abundance and body size.

In this paper we model the process of taxonomic evolution as a Galton-Watson branching process in discrete time and, using maximum likelihood, develop methods to estimate the probabilities of origination, persistence, and extinction of fossil taxa. We use the methods to estimate the probabilities of origination, persistence, and extinction of families (1) within 135 orders of marine invertebrate organisms, (2) within 12 phyla, and (3) within all marine invertebrate life (independently of the suprafamilial classification).

Most orders, including the arcoid bivalves, the dentaloid scaphopods, the orders of chitins, and many others, have relatively low probabilities of familial origination and extinction. The various ammonoid and trilobite orders, and some others, have high probabilities of origination and extinction. Among the phyla, the Archaeocyatha have the highest probabilities of familial origination and extinction, and the Annelida the lowest, with the more typical phyla of shelly organisms clustering near the high end of the probability scale. The Porifera and Protozoa also have low probabilities but not as low as the Annelida. The estimated origination and extinction probabilities for families within all marine invertebrate life are 0.470 and 0.452 per stage, respectively, values that are at the high end of the probability scale. We have also estimated the probabilities of ultimate extinction (extinction of all families) of the supertaxa.

By analyzing the changes of the diversity during each stratigraphic stage separately, we have also determined the trajectories of the estimated origination and extinction probabilities for families within all marine invertebrate life. The estimated origination probability is relatively high in association with the expansion of the Cambrian and Paleozoic evolutionary faunas and declines to more normal levels for the remainder of the Phanerozoic. The trajectory of the estimated extinction probability is from nearly zero early in the Phanerozoic to more normal levels later, showing clearly defined peaks in association with the five Phanerozoic mass-extinction events. The terminal Cretaceous mass extinction is the only one of the five that was not preceded by a monotonic decline of origination probability or by a series of stages with low origination probability. It appears to have been a unique, singular event.

Because the mathematical theory we employ as a model corresponds so closely to the processes of taxonomic evolution as we understand them, we believe that the theory provides a reasonable model of biological reality.

The Late Neogene vertebrate fossil record from Yushe Basin presents multiple, superposed assemblages from a single area, spanning roughly the interval of 6–2 Ma. Both large and small mammals show peak species richness in the middle Pliocene but indicate relative faunal stability throughout the Pliocene. Large mammals show turnover, especially extinction, around 5 and 2.5 Ma. Small mammals indicate change (over half of the species and several genera), as well as turnover at the species level, between 4 and 3.4 Ma. The loosely controlled dating of these events does not disprove hypothetical correlation with events in North America and with global climatic shifts. Elements that lack Yushe antecedents, some being long-distance dispersers, appear throughout the section, but with little effect on the resident assemblage. First records of well-documented immigrants (from North America, Europe, Africa, southern Asia, or high latitudes) generally do not coincide with ecomorph extinctions. Early Pliocene exchange between Asia and North America appears to have been balanced in both directions and involved a small proportion of the fauna. Immigration probably was opportunistic and contributed to faunal enrichment. We interpret the Yushe Pliocene mammalian assemblages as representing a fauna that was stable from ca. 5 to 2.5 Ma and changed mainly by additions and congeneric species substitutions.

Onshore-offshore patterns of faunal change occurred at many taxonomic scales during the Paleozoic Era, ranging from replacement of the Cambrian evolutionary fauna by the Paleozoic fauna to the environmental expansion of many orders and classes. A simple mathematical model is constructed to investigate such change. The environmental gradient across the marine shelf-slope is treated as a linear array of discrete habitats, each of which holds a set number of species, as observed in the fossil record. During any interval of time, some portion of the species in each habitat becomes extinct by background processes, with rates of extinction varying among both clades and habitats, as also observed in the record. After extinction, species are replaced from within the habitat and from immediately adjacent habitats, with proportions dependent on surviving species. This model leads to the prediction that extinction-resistant clades will always diversify at the expense of extinction-prone clades. But if extinction intensity is highest in nearshore habitats, extinction-resistant clades will expand preferentially in the onshore direction, build up diversity there, and then diversify outward toward the offshore. Thus, onshore-offshore patterns of diversification may be the expectation for faunal change quite independently of whether or not clades originate onshore. When the model is parameterized for Paleozoic trilobites and brachiopods, numerical solutions exhibit both a pattern of faunal change and a time span for diversification similar to that seen in the fossil record. They also generate structure similar to that seen in global diversification, including logistic patterns of growth, declining origination but constant extinction within clades through time, and declining overall extinction across clades through time.

The depositional history of Upper Miocene through Recent sediments from DSDP Site 214 (Ninetyeast Ridge, Indian Ocean) is reexamined. Samples of the Globorotalia tumida planktic foraminiferal lineage, originally obtained from these sediments by Malmgren et al. (1983), serve as the empirical basis for the recognition of punctuated anagenesis as a distinct mode of phenotypic evolution and have been the subject of numerous additional investigations. However, conclusions reached by previous authors depend strictly on the validity of the original chronostratigraphic interpretation of these sediments. Graphic correlation analysis of first- and last-appearance datum levels for a total of 41 planktic foraminiferal, radiolarian, and calcareous nannoplankton taxa provides evidence for a more complex depositional history at this deep-sea site than originally believed. Based on a conservative model of variation in the pattern of sediment accumulation rates, the lowermost portion of the studied section (6.5-4.3 Ma) represents an interval of temporally condensed sediment accumulation (1.88 cm/1,000 yr) followed by an interval (4.3-2.8 Ma) of temporally expanded sediment accumulation (3.97 cm/1,000 yr). This interval, in turn, is followed by a depositional hiatus or an extremely condensed interval, at least 800,000 yr in duration, which is followed by another relatively condensed (1.36 cm/1,000 yr) interval from 2.0 Ma-Recent. Although this chronostratigraphic reinterpretation deviates substantially from the original, which recognized Site 214 as being both temporally continuous and exhibiting a constant sediment accumulation rate from the Upper Miocene through the Upper Pliocene, it is more consistent with expectations based on Neogene eustatic sea-level fluctuations and global surveys of Neogene hiatus distributions. Age assignments for samples of the Gr. tumida lineage based on the revised chronostratigraphic model reverse some findings of previous investigators with respect to the distinctiveness of phenotypic evolutionary rates characterizing the transition from Gr. plesiotumida to Gr. tumida. Finally, a brief survey of similar marine invertebrate lineage studies shows that changes in the rate of phenotypic evolution often appear to coincide with major physical changes in the paleoceanographic environment. Such correspondences may be due, at least in part, to the effect of these environmental changes on sediment accumulation rates. Paleobiologists who seek to understand patterns of phenotypic change over time must remove the effects of variations in sediment accumulation rates from their data before evolutionary hypothesis testing and remain aware of the limitations imposed on their interpretations by the uncertain nature of chronostratigraphic inference.

The directionality of long-term trends can be strongly biased by forces intrinsic to a clade. Trends in body size and skeletal shape may be dictated more by variations in survivorship that reflect differences in ecology than by long-term directional changes in the environment. Hence, mass extinctions can help drive evolutionary trends by selectively eliminating some morphologies and permitting the survivors to found the next radiation.

Examples include repeated trends toward larger maximum body size and the evolution of keeled species from those with globose tests in planktonic foraminifera. Both the trends in size and shape develop because small species with globose tests are significantly more resistant to extinction than species that are large or have peripheral keels. Hence, the survivors of both the Cretaceous-Tertiary and Eocene-Oligocene extinction episodes are small, unkeeled taxa. Large species and species with keels evolved convergently after both mass extinctions as the founders radiated anew.

Comparison of three radiations of planktonic foraminifera suggest that the convergent evolution of similar test shapes and sizes is not due to synchronous changes in oceanography that track evolutionary trends. Instead, the reestablishment of habitat heterogeneity is needed to permit the ensuing radiation to unfold rather than to closely guide its progress. Similar evolutionary trends will develop in each radiation as long as the founders have similar morphology and the evolution of variants present in the previous radiation is not precluded by the environment.