In both jaw geometry and molar morphology, eutherian carnivores (order Carnivora) as a whole display greater diversity (plasticity in evolution from the primitive type) than marsupial carnivores (order Dasyurida). This is related to the difference in tooth replacement between the two taxa. In Carnivora, the permanent carnassial is preceded by a deciduous carnassial; the permanent tooth can erupt in its (geometrically) permanent position, and the post-carnassial molars are free to evolve for specialized functions or be reduced. In Dasyurida, there is relative molar progression, each erupting molar in turn functioning as a carnassial, and subsequently being pushed forwards in the jaw by the next erupting molar. Thus, all molars have carnassiform morphology, and none are free to develop for other functions. The greater plasticity of Carnivora has led to their adaptive zone being broader (as a group they are relatively more eurytopic than Dasyurida), which in turn has led to greater taxonomic diversity within Carnivora than Dasyurida. The resulting pattern from a macroevolutionary point of view is that, even in the absence of direct competition, Carnivora have had greater evolutionary “success” than Dasyurida.

Analysis of post-mortem buoyancy loss in Nautilus shells suggests that extensive nekroplanktonic drifting occurs infrequently. Most shells do not reach the surface but settle to the sea floor, after a short period of ascent. This occurs because the rate of water influx into the phragmocone due to ambient hydrostatic pressure is sufficiently rapid in most cases to overcome positive buoyancy before the shell reaches the surface. The resulting geographic distribution of Nautilus shells would therefore mirror the distribution of the live animals. Thus, post-mortem drift in Nautilus cannot be used as a basis for questioning the validity of cephalopod paleobiogeography. Estimate of influx rates in ammonoid siphuncles indicates that many, if not most, ammonoid shells also would not become nekroplanktonic. This is especially true for small (<5 cm diameter) shells. Cephalopod paleobiogeographic investigation appears less subject to criticism stemming from the supposed obfuscating effects of post-mortem drift than previously thought.

Paleoecological analyses that test for spatial or temporal variation in diversity must consider not only sampling and preservation bias, but also the effects of temporal scale (i.e., time-averaging). The species-time relationship (STR) describes how species diversity increases with the elapsed time of observation, but its consequences for assessing the effects of time-averaging on diversity of fossil assemblages remain poorly explored. Here, we use a neutral, dispersal-limited model of metacommunity dynamics, with parameters estimated from living assemblages of 31 molluscan data sets, to model the effects of within-habitat time-averaging on the mean composition and multivariate dispersion of assemblages, on diversity at point (single station) and habitat scales (pooled multiple stations), and on beta diversity. We hold sample size constant in STRs to isolate the effects of time-averaging from sampling effects. With increasing within-habitat time-averaging, stochastic switching in the identity of species in living (dispersal-limited) assemblages (1) decreases the proportional abundance of abundant species, reducing the steepness of the rank-abundance distribution, and (2) increases the proportional richness of rare, temporally short-lived species that immigrate from the neutral metacommunity with many rare species. These two effects together (1) can shift the mean composition away from the non-averaged (dispersal-limited) assemblages toward averaged assemblages that are less limited by dispersal, resembling that of the metacommunity; (2) allow the point and habitat diversity to increase toward metacommunity diversity under a given sample size (i.e., the diversity in averaged assemblages is inflated relative to non-averaged assemblages); and (3) reduce beta diversity because species unique to individual stations become shared by other stations when limited by a larger but static species pool. Surprisingly, these scale-dependent changes occur at fixed sample sizes and can become significant after only a few decades or centuries of time-averaging, and are accomplished without invoking ecological succession, environmental changes, or selective postmortem preservation. Time-averaging results in less inflation of diversity at habitat than at point scales; paleoecological studies should thus analyze data at multiple spatial scales, including that of the habitat where multiple bulk samples have been pooled in order to minimize time-averaging effects. The diversity of assemblages that have accumulated over 1000 years at point and habitat scales is expected to be inflated by an average of 2.1 and 1.6, respectively. This degree of inflation is slightly higher than that observed in molluscan death assemblages at these same spatial scales (1.8 and 1.3). Thus, neutral metacommunity models provide useful quantitative constraints on directional but predictable effects of time-averaging. They provide minimal estimates for the rate of increase in diversity with time-averaging because they assume no change in environmental conditions and in the composition of the metacommunity within the window of averaging.

Physical environmental factors have been seen as paramount in determining many large-scale biodistributional patterns in time and space. Although this is probably correct for many situations, this view has become so pervasive that it has led to the neglect of the role of biotic interactions in setting large-scale diversity patterns. (In this paper diversity denotes taxonomic richness.) New approaches to this perennial debate on the roles of physical and biotic forces in paleoecology and macroevolution are needed, and here we explore an argument for the role of incumbency or priority effects in the dynamics behind the most dramatic spatial pattern in biodiversity, the latitudinal diversity gradient.

Although available paleobiological data indicate that the geographic ranges of marine species are maintained throughout their entire observable durations, other evidence suggests, by contrast, that the ranges of higher taxa expand as they age, perhaps in association with increased species richness. Here, I utilize a database of Ordovician genus occurrences collected from the literature for several paleocontinents to demonstrate that a significant aging of the global biota during the Ordovician Radiation was accompanied by a geographic and environmental expansion of genus ranges. The proportion of genera occurring in two or more paleocontinents in the database, and two or more environmental zones within a six-zone onshore-offshore framework, increased significantly in the Caradocian and Ashgillian. Moreover, widespread genera tended to be significantly older than their endemic counterparts, suggesting a direct link between their ages and their environmental and geographic extents. Expansion in association with aging was corroborated further by demonstrating this pattern directly among genera that ranged from the Tremadocian through the Ashgillian. Taken together, these results are significant not only for what they reveal about the kinetics of a major, global-scale diversification, but also for what they suggest about the interpretation of relationships between diversity trends at the α (within-community) and β (between-community) levels.

When assessing the timing of branching events in a phylogeny, the most important tools currently recognized are a reliable molecular phylogeny and a continuous, relatively complete fossil record. Coralline algae (Rhodophyta, Corallinales, and Sporolithales) constitute an ideal group for this endeavor because of their excellent fossil record and their consistent phylogenetic reconstructions. We present the evolutionary history of the corallines following a novel, combined approach using their fossil record, molecular phylogeny (based on the 18S rDNA gene sequences of 39 coralline species), and molecular clocks. The order of appearance of the major monophyletic taxa of corallines in the fossil record perfectly matches the sequence of branching events in the phylogeny. We were able to demonstrate the robustness of the node ages in the phylogeny based on molecular clocks by performing an analysis of confidence intervals and maximum temporal ranges of three monophyletic groups of corallines (the families Sporolithaceae and Hapalidiaceae, as well as the subfamily Lithophylloideae). The results demonstrate that their first occurrences are close to their observed appearances, a clear indicator of a very complete stratigraphic record. These chronological data are used to confidently constrain the ages of the remaining branching events in the phylogeny using molecular clocks.

Ecomorphological diversity of Mesozoic mammals was presumably constrained by selective pressures imposed by contemporary vertebrates. In accordance, Mesozoic mammals for a long time had been viewed as generalized, terrestrial, small-bodied forms with limited locomotor specializations. Recent discoveries of Mesozoic mammal skeletons with distinctive postcranial morphologies have challenged this hypothesis. However, ecomorphological analyses of these new postcrania have focused on a single taxon, a limited region of the skeleton, or have been largely qualitative.

For more comprehensive locomotor inference in Mesozoic mammals, we applied multivariate analyses to a morphometric data set of extant small-bodied mammals. We used 30 osteological indices derived from linear measurements of appendicular skeletons of 107 extant taxa that sample 15 orders and eight locomotor modes. Canonical variate analyses show that extant small-bodied mammals of different locomotor modes have detectable and predictable morphologies. The resulting morphospace occupation reveals a morphofunctional continuum that extends from terrestrial to scansorial, arboreal, and gliding modes, reflecting an increasingly slender postcranial skeleton with longer limb output levers adapted for speed and agility, and extends from terrestrial to semiaquatic/semifossorial and fossorial modes, reflecting an increasingly robust postcranial skeleton with shorter limb output levers adapted for powerful, propulsive strokes. We used this morphometric data set to predict locomotor mode in ten Mesozoic mammals within the Docodonta, Multituberculata, Eutriconodonta, “Symmetrodonta,” and Eutheria. Our results indicate that these fossil taxa represent five of eight locomotor modes used to classify extant taxa in this study, in some cases confirming and in other cases differing from prior ecomorphological assessments. Together with previous locomotor inferences of 19 additional taxa, these results show that by the Late Jurassic mammals had diversified into all but the saltatorial and active flight locomotor modes, and that this diversification was greatest in the Eutriconodonta and Multituberculata, although sampling of postcranial skeletons remains uneven across taxa and through time.

Macroevolutionary change from the Middle to the Late Paleozoic crinoid fauna was not the result of mass extinction. The presumption that the decline of the middle Paleozoic crinoid fauna was from a single mass extinction event was tested using seriation, multidimensional scaling (MDS), binomial analysis, and bootstrapping simulations on a data set which is a comprehensive revision of old faunal lists. The data for these analyses were based on temporal distributions of 214 species from 69 late Osagean and early Meramecian localities from the midcontinental United States. The time under consideration is subdivided into seven informal intervals using MDS in conjunction with biostratigraphy. Seriation of species ranges into these intervals results in a gradual pattern of faunal turnover, and sampling bias can be eliminated as a cause for this more gradual pattern. MDS analysis of the crinoid range data is similar to MDS simulations using data with continuous, monotonic species turnover and dissimilar to a simulated mass extinction. Binomial analysis and bootstrapping demonstrate that the observed number of extinctions at the putative extinction boundary were not unusually high. All methods agree that extinctions throughout this time were high but spanned several time intervals and that rapid, monotonic faunal turnover describes the data better than mass extinction. Macroevolutionary processes other than mass extinction and microevolutionary processes must have dictated the character and composition of this remarkable faunal transition among the Crinoidea.

Like living herbivorous lizards, chelonians, birds, and mammals, plant-eating dinosaurs probably relied on a symbiotic gut microflora, housed in a hindgut fermentation chamber, to break down plant cell wall constituents. Large body sizes in most herbivorous dinosaurs resulted in low mass-specific metabolic rates and low rates of digesta passage through the gut; the effects of large body size were probably enhanced by the low metabolic rates of large dinosaurs as compared with large mammals. The long residence time of digesta in the gut permitted long exposure of refractory plant materials to the microflora, probably enabling even those dinosaurs with unsophisticated dentitions to survive on fodder with high fiber content. Large herbivorous dinosaurs probably fed on plants whose allelochemical defenses were geared more toward reducing digestibility than attacking the herbivore's metabolism directly, obviating the need for a foregut fermentation chamber and permitting these large herbivores to take advantage of the energetic benefits of hindgut fermentation for digestion of low-quality fodder. Differences in dentitions among the groups of herbivorous dinosaurs may correlate with differences in standard metabolic rate, activity level, body size, or food quality, or combinations of these factors, but the relative importance of each is difficult to assess. Because the mass of the fermentation contents was probably large in big herbivorous dinosaurs, the heat of fermentation may have been a significant source of thermoregulatory heat for these reptiles.

Both intensive analysis of Gryphaea and an extensive survey of Jurassic bivalves in Europe supports the punctuated equilibria model of speciation, with the exception of phyletic size increase, which appears within the limits of the data to be gradualistic and affects at least a significant minority of species. The size increase takes place both within species and between successive species in a lineage. In Liassic Gryphaea, a combination of allometry in the ontogenetic development of the ancestral species and paedomorphosis led also to changes of shape up the sequence. The production of evolutionary trends by species selection is not supported.

The favoured interpretation of phyletic size increase is a gradual delay in maturation time consequent upon a change in the organisms' adaptive strategy from the r-selected to the K-selected mode and is backed up by evidence on changing population numbers and distribution. An extinction/speciation model is proposed for neritic organisms based on fluctuating sea levels. Times of low sea level or regression correspond with times of high stress and hence high extinction among stenotopic organisms and increased rates of allopatric speciation, with r-selection as the dominant mode. Times of high sea level or transgression correspond with low extinction and speciation rates and increased freedom of migration. K-selection is the dominant mode and often leads to phyletic size increase provided the environment remains stable for a sufficiently long period.

This summer's meeting season saw five gatherings devoted specifically to the topic of extinction: Chicago's Field Museum symposium in May (see Diamond 1983); a field conference to examine the Cretaceous-Tertiary transition in Montana; a symposium on molluscan extinction at the American Malacological Union meeting in Seattle; a special session at the annual meeting of the Society for the Study of Evolution in St. Louis; and a research conference on the campus of Northern Arizona University in Flagstaff (see also Lewin 1983a,b,c). Several recent books consider the topic (Ehrlich and Ehrlich 1981; Nitecki 1981; Silver and Schultz 1982; Martin and Klein, 1984) and plans are underway to publish the papers presented at the Field Museum, Seattle, and Flagstaff meetings. Our review of the topic is based principally on the themes explored at the St. Louis and Flagstaff meetings.

Size distributions, based on molar length, are presented for various groups of living and fossil primates. I examine (1) the extent to which particular taxonomic groups with distinctive morphological and behavioral attributes are characterized by distinctive size ranges or distributions and (2) the way in which size ranges and distributions are affected by the presence or absence of other primates within the same geographical area and time. Distinctive behavioral and ecological “adaptive zones” are characterized by distinctive size ranges and behavioral and morphological parallelism or convergence among living and fossil primate taxa usually results in similar size distributions.

Biotic invasions are a common feature of the fossil record, yet remarkably little is known about them, given their enormous potential to reveal the processes that regulate local and regional diversity over long time scales. We used additive diversity partitioning to examine how diversity structure changed as a result of a marine biotic invasion in tropical, shallow and deep subtidal environments spanning approximately 4 Myr in the Late Ordovician. The biotic invasion increased richness in the regional ecosystem by nearly 40%. Within-habitat turnover diversity accounts for most of the increase in richness, with between-habitat turnover diversity contributing a lesser amount. Increases in these components of diversity were accommodated by increased packing of species along a depth gradient and increased habitat heterogeneity. Diversity metrics that incorporate taxon abundance (Shannon information, Simpson's D) show similar patterns and reveal that many invading taxa were locally abundant and widespread in their occurrence. Extinction of incumbent taxa did not foster the invasion; rather the invasion appears to be linked to a regional or global warming event. Taken together, these observations indicate that these Late Ordovician marine communities were open to invasion and not saturated with species. Moreover, the increase in species diversity caused by the invasion was not ephemeral; instead it lasted for at least 1 Myr. Similar studies of other biotic invasions in the fossil record are necessary to determine (1) the factors, such as extinction of incumbents or resource limitation, that may facilitate or inhibit invasion in ancient ecosystems; (2) how local and regional ecosystems respond to invasion; and (3) the extent to which biotic invasions play a substantial role in ecosystem change through geologic time.

Marsupial mammals are characterized by a pattern of dental replacement thought to be unique. The apparent primitive therian pattern is two functional generations of teeth at the incisor, canine, and premolar loci, and a series of molar teeth, which by definition are never replaced. In marsupials, the incisor, canine, and first and second premolar positions possess only a single functional generation. Recently this pattern of dental development has been hypothesized to be a synapomorphy of metatherians, and has been used to diagnose taxa in the fossil record. Further, the suppression of the first generation of teeth has been linked to the marsupial mode of reproduction, through the mechanical suppression of odontogenesis during the period of fixation of marsupials, and has been used to reconstruct the mode of reproduction of fossil organisms. Here we show that dental development occurs throughout the period of fixation; therefore, the hypothesis that odontogenesis is mechanically suppressed during this period is refuted. Further, we present comparative data on dental replacement in eutherians and demonstrate that suppression of tooth replacement is fairly common in diverse groups of placental mammals. We conclude that reproductive mode is neither a necessary nor a sufficient explanation for the loss of tooth replacement in marsupials. We explore possible alternative explanations for the loss of replacement in therians, but we argue that no single hypothesis is adequate to explain the full range of observed patterns.

Barstovian (medial Miocene) mammalian faunas from the Texas Gulf Coastal Plain contained four apparently sympatric species of rhinoceroses: the common forms Aphelops megalodus and Teleoceras medicornutus, a dwarf Teleoceras, and a dwarf Peraceras. Previous work has suggested positive allometry in tooth area with respect to body size in several groups of mammals, i.e., larger mammals have relatively more tooth area. However, dwarfing lineages were shown to have relatively more tooth area for their body size. Our data show no significant allometry in post-canine tooth area of either artiodactyls or ceratomorphs. Similarly, dwarf rhinoceroses and hippopotami show no more tooth area than would be predicted for their size. Limbs are proportionately longer and more robust in larger living ceratomorphs (rhinos and tapirs) than predicted by previous authors. Limb proportions of both dwarf rhinoceroses and dwarf hippopotami are even more robust than in their living relatives.

The high rhinoceros diversity reflects the overall high diversity of Barstovian faunas from the Texas Gulf Coastal Plain. The first appearance of several High Plains mammals in these faunas indicates “ecotone”-like conditions as faunal composition changed. Study of living continental dwarfs shows that there is commonly an ecological separation between browsing forest dwarfs and their larger forebears, which are frequently savannah grazers. This suggests that the dwarf rhinoceroses might have been forest browsers which were sympatric with the larger grazing rhinos of the High Plains during the Barstovian invasion. The continental dwarf model also suggests that insular dwarfism may be explained by the browsing food resources that predominate on islands.

In the past decade paleobiologists have applied the techniques of both ecological and historical biogeography, although vicariance/cladistic approaches have as yet had minimal impact. The traditional focus of paleobiogeographic study has been the province, a statistical entity defined by clusters of range endpoints of individual taxa. The study of such provinces has been useful in inferring past continental positions (although ambiguities remain that must be resolved using independent geological criteria) and in understanding the role of past global geographies in regulating biotic diversity through changes in the numbers and extent of provinces. This approach can be complemented by the treatment of geographic ranges of taxa as irreducible or emergent traits with far-reaching evolutionary effects upward and downward within a genealogical hierarchy. Temperature tolerances in benthic marine organisms appear to be by-products of selection for enzyme structures imparting favorable activity levels within the normal temperature range rather than direct products of selection for resistance to temperature extremes. Thus geographic range endpoints, which are also influenced by dispersal capability and the resulting scale of gene flow among disjunct populations, are not direct products of selection. However, the magnitudes of geographic ranges of species and clades behave as emergent properties and significantly influence taxonomic survivorship during background and mass extinctions in ways that are not extrapolations of effects at lower hierarchical levels. Biogeography shapes macroevolutionary patterns of origination and extinction during times of normal, background extinction and mass extinction. Preferential extinction among regions or among endemic rather than widespread clades can result in strong biases in the nature of the survivors of mass extinctions, with taxa being lost not because of selection against attributes of individual organisms but because of higher-order patterns of geographic selectivity.

We compare the preservation (taphonomic grade) and age of Chione (bivalve) and foraminifera from modern siliciclastic tidal flat sediments of Bahia la Choya, Sonora, Mexico (northern Gulf of California). Disarticulated shells of Chione collected from the sediment-water interface of Choya Bay exhibit a substantial range in taphonomic grade and age, several hundred years to ∼80–125 ka based on Accelerator Mass Spectrometer 14C dates and D-Alloisoleucine/L-Isoleucine values. There is not, however, a one-to-one correspondence between age and taphonomic alteration of Chione: old (or young) valves may be highly altered or they may be relatively pristine. In contrast to Chione, most foraminiferal tests at Choya Bay are quite pristine, which suggests a quite young age, but tests are surprisingly old (up to ∼2,000 calendar years based on Accelerator Mass Spectrometer 14C dates).

We suggest that following seasonal pulses in reproduction, some foraminiferal tests are rapidly incorporated into a subsurface shell layer by “Conveyor Belt” deposit feeders and preserved there, while the rest of the reproductive pulse rapidly dissolves. Ultimately, some of these buried tests, along with Chione, are transported back to the surface by biological activity and storms. The much greater range of taphonomic grades and ages among Chione shells suggests that they, unlike foraminifera, are sufficiently large and preservable (low surface/volume ratio and chemical reactivity) to undergo many cycles of degradation, burial, and exhumation before complete destruction. The age of foraminiferal tests indicates that time-averaging of microfossil assemblages at Choya Bay is much more insidious than would be expected considering the relatively pristine state of the tests alone.

Based on our studies, the lower limit of temporal resolution of shallow shelf microfossil assemblages appears to be ∼1000 years. We caution, however, that each depositional setting (taphofacies) should be evaluated on a case-by-case basis before gross generalizations are made. Indeed, the discrepancy between age and taphonomic grade of fossil assemblages at Choya Bay suggests that neither hardpart size or taphonomic grade are infallible indicators of test preservability or likely temporal resolution of the host assemblage, and that the dynamics of hardpart input and loss must also be evaluated.

Geographic range and taxonomic duration are known to be positively correlated in a number of biologic groups; this is usually attributed to the influence of range upon duration rather than the other way about. Here we analyze two distinct components of this correlation within species and genera of marine invertebrates and microfossils by partitioning the total duration into two parts: the time it takes a taxon to attain its maximum geographic range, and the time a taxon persists after attaining its peak range. We find that the longer it takes a taxon to attain its maximum geographic range, the wider is that range. We also find that the broader the maximum range, the greater is the duration after this maximum is attained. These two correlations are equally strong on average. There is thus a reciprocal relationship between duration and geographic range, and there is no compelling evidence that range generally determines duration more or less than duration determines range.

The problem of gradual versus punctuated change within phyletic lineages can be understood in terms of the homogeneity of evolutionary dynamics. Hypotheses of punctuated change imply that the rules governing evolutionary change shift over time such that the normal dynamics of stasis are temporarily suspended, permitting a period of net evolutionary change. Such explanations are members of a larger class of models in which evolutionary dynamics are in some way heterogeneous over time. In this paper, I develop a likelihood-based statistical framework to evaluate the support for this kind of evolutionary model. This approach divides evolutionary sequences into nonoverlapping segments, each of which is fit to a separate evolutionary model. Models with heterogeneous dynamics are generally more complex—they require more parameters to specify—than uniform evolutionary models such as random walks and stasis. The Akaike Information Criterion can be used to judge whether the greater complexity of punctuational models is offset by a sufficient gain in log-likelihood for these models to be preferred.

I use this approach to analyze three case studies for which punctuational explanations have been proposed. In the first, a model of punctuated evolution best accounted for changes in pygidial morphology within a lineage of the trilobite Flexicalymene, but the uniform model of an unbiased random walk remains a plausible alternative. Body size evolution in the radiolarian Pseudocubus vema was neither purely gradual nor completely pulsed. Instead, the best-supported explanation posited a single, pulsed increase, followed later by a shift to an unbiased random walk. Finally, for the much-analyzed claim of “punctuated gradualism“ in the foraminifera Globorotalia, the best-supported model implied two periods of stasis separated by a period of elevated but not inherently directional evolution. Although the conclusions supported by these analyses generally refined rather than overturned previous views, the present approach differs from those prior in that all competing interpretations were formalized into explicit statistical models, allowing their relative support to be unambiguously compared.

Sauropod dinosaurs were the largest terrestrial animals and their growth rates remain a subject of debate. By counting growth lines in histologic sections and relating bone length to body mass, it has been estimated that Apatosaurus attained its adult body mass of about 25,000 kg in as little as 15 years, with a maximum growth rate over 5000 kg/yr. This rate exceeds that projected for a precocial bird or eutherian mammal of comparable estimated body mass. An alternative method of estimating limb length and body mass for each growth line, and fitting the resulting age/mass data to the von Bertalanffy growth equation, yields a revised growth curve suggesting that Apatosaurus adult mass was reached by 70 years with a maximum growth rate of 520 kg/yr. This alternative method for growth rate determination can also be applied to histological studies of other sauropods. At only about half the mass of Apatosaurus, Janenschia took between 20 and 30 years to attain its adult size (over 14,000 kg). This result is supported by independent evidence of estimated bone apposition rates. Despite having an adult body mass greater than Apatosaurus, the titanosaurid Alamosaurus attained a mass over 32,000 kg within 45 years and a maximum growth rate of 1000 kg/yr. Titanosaurids may have been the fastest growing of all sauropods. Even so, sauropod growth rate estimates produced using the von Bertalanffy equation fall between those projected for reptiles and those for precocial birds of equivalent projected body mass. These results are comparable to those found for smaller dinosaurs, and suggest that sauropods grew at rates similar to other dinosaurs in spite of their great size.