One of the striking yet scarcely documented episodes of clade replacement in the post-Paleozoic fossil record is the decline of cyclostome Bryozoa and the corresponding, rapid diversification of cheilostome Bryozoa. These clades are closely associated morphologically and phylogenetically, and their ecological similarities have previously led to the inference that competition was a primary cause of the overt pattern of replacement. Alternatively, previous compilations of bryozoan families and genera have implied that extinctions at the Cretaceous/Tertiary boundary differentially affected cyclostomes, and thus were also an important factor in the transition.

We first evaluated the ecological context for competition between the two clades, then updated and reexamined the history of absolute family diversity for bryozoans in consecutive geologic stages from Jurassic to Recent. The resulting trends echo the patterns shown in earlier family level compilations, but indicate a slight shift in the frequency of cheilostome family originations from Late Cretaceous to early Paleogene. The relative fall in cyclostome family diversity at the Cretaceous/Tertiary boundary is significantly less than shown in earlier genus level compilations. We then assessed these various compilations of absolute diversity by analyzing species counts and percentages in 728 fossil assemblages, primarily from North America and Europe, over the same time interval. Cyclostome species overwhelmingly dominate assemblages from Jurassic through Cenomanian, then decline significantly in average percentage dominance through the Campanian. Cheilostomes are predominant in Campanian and later assemblages. Cyclostome species percentages do decrease overall through the Tertiary, but this decrease is small and non-uniform, varying around 30%, with a sharp drop in the Late Neogene. Our within-assemblage results indicate that as cheilostomes radiate, their mean species diversity, maximum diversity, and variance all increase, thereby accounting for much of the decline in average percentage of cyclostomes within assemblages. While this result does not exclude a role for competition, an hypothesis of relative decline in cyclostome species richness based on competitive extinction alone seems unlikely. Further, despite decreases in absolute species counts following end-Cretaceous extinctions, within-assemblage percentages of cheilostome or cyclostome species show only slight change relative to one another. Comparison of these and earlier diversity compilations indicates that the dynamics of bryozoan clade replacement may be perceived differently at different ecologic scales or taxonomic ranks.

More than 5000 measurements were taken on over 1000 specimens of two species of brachiopods, Mediospirifer audaculus and Athyris spiriferoides, from the Middle Devonian Hamilton Group of New York state. Statistical analyses were performed on these data, with specimens partitioned by their occurrence in one of many paleoenvironments and stratigraphic horizons. Neither species showed substantial morphological departures between first appearance and extinction (the range of the Hamilton Group, roughly 5 m.y.). However, oscillations in morphology were discovered in both taxa.

For the two species we studied, groups of organisms occurring in a single paleoenvironment undergo moderate morphological change through time; however, the net sum of changes through time in all paleoenvironments in which these species occur is essentially zero. Therefore, stasis may be partly a property of the organization of species into different environmental populations. Different “environmental populations” may evolve, but they will typically do so in several different “directions,” generally producing no net change. The difference between the morphology of species in different environments over the whole interval of the Hamilton Group is also nil, thereby ruling out any major role that ecophenotypic effects could play in the patterns recognized herein.

Biological veracity of the sharp diversity increase observed in many analyses of the post-Paleozoic marine fossil record has been debated vigorously in recent years. To assess this question for sample-level (“alpha”) diversity, we used bulk samples of shelly invertebrates, representing three major fossil groups (brachiopods, bivalves, and gastropods), to compare the Jurassic and late Cenozoic sample-level diversity of marine benthos. After restricting the data set to single-bed, whole-fauna, bulk samples (n ≥ 30 specimens) from comparable open marine siliciclastic facies, we were able to retain 427 samples (255 Jurassic and 172 late Cenozoic), with most of those samples originating from our own empirical work.

Regardless of the diversity metric applied, the initial results suggest that standardized sample-level species (or genus) diversity, driven by evenness and/or richness of the most common taxa, increased between the Jurassic and late Cenozoic by at least a factor of 1.6. When the data are partitioned into the three dominant higher taxa, it becomes clear that (1) the bivalves, which dominated the samples for both time intervals, increased in sample-level diversity between the Jurassic and the late Cenozoic by a much smaller factor than the total fauna; (2) the removal of brachiopods, which were a noticeable component of the Jurassic samples, did not significantly affect standardized sample-level diversity estimates; and (3) the gastropods, which were rare in the Jurassic but common in many late Cenozoic samples, contributed notably to the increase in sample-level diversity observed between the two time intervals. Parallel to these changes, the samples revealed secular trends in ecological structure, including Jurassic to late Cenozoic increases in proportion of (1) infauna, (2) mobile forms, and (3) non-suspension-feeding organisms. These trends mostly persist when data are restricted to bivalves.

Supplementary analyses indicate that these patterns cannot be attributed to sampling heterogeneities in paleolatitudinal range, lithology, or paleoenvironment of deposition. Likewise, when data are restricted to samples dominated by species with originally aragonitic shells, the observed temporal changes persist at a comparable magnitude, suggesting that the pervasive loss of aragonite in the older fossil record is unlikely to have been the primary cause of the observed patterns. The comparable ratio of identified to unidentified species and genera, observed when comparing the Jurassic and late Cenozoic samples, indicates that the relatively poorer (mold/cast) preservation of Jurassic aragonite species also is unlikely to have been responsible for the observed patterns. However, the diagenesis-related taphonomic and methodological artifacts cannot be ruled out as an at least partial contributor to the observed post-Paleozoic changes in diversity, taxonomic composition, and ecology (the outcomes of the three tests of the diagenetic bias available to us are incongruent).

The study demonstrates that the post-Paleozoic trends in the sample-level diversity, ecology, and taxonomic structure of common taxa can be replicated across multiple studies. However, the diversity increase estimated here is much less prominent than suggested by many previous analyses. The results also narrow the list of causative explanations down to two testable hypotheses. The first is diagenetic bias—a spurious trend driven by either (a) increasing taphonomic loss of small specimens in the older fossil record or (b) a shift in sampling procedures between predominantly lithified rocks of the Mesozoic and predominately unlithified, and therefore sievable, sediments of the late Cenozoic. The second hypothesis is genuine biological changes—macroevolutionary trends in the structure of marine benthic associations through time, consistent with predictions of several related models such as evolutionary escalation, increased ecospace utilization, and the Mesozoic marine revolution. Future studies should focus on testing these two rival models, a key remaining challenge for identifying the primary causative mechanism for the long-term changes in sample-level diversity, ecology, and taxonomic structure observed in the Phanerozoic marine fossil record.

A compilation of species occurrences in a chronostratigraphic framework of depositional sequences from a 250,000 km2 area in the eastern United States is used to test for coordinated stasis in Middle and Upper Ordovician articulate brachiopods. Two rapid pulses of turnover in brachiopod species separate three periods of relatively lower turnover (ecologic-evolutionary [EE] subunits) that range from 3 to 9 m.y. in duration. Turnover within these EE subunits is characterized by high levels of percent species origination (ca. 60%) and percent species extinction (ca. 80%) and low levels of percent species persistence (<10%), all of which fall outside the range reported for coordinated stasis. Turnover between EE subunits is characterized by low levels of percent species holdover and percent species carryover (ca. 10% or less) and is consistent with coordinated stasis, although turnover pulses are driven largely by pulses in either extinction or origination, and not by pulses in both, as reported for coordinated stasis. Taken together, although these data display a marked bimodality in turnover, high levels of turnover within EE subunits is inconsistent with a pattern of coordinated stasis. Turnover rates within these EE subunits are much higher than previous global estimates for Cambro-Ordovician brachiopods or Phanerozoic marine species and indicate that local extirpation and migration play a significant role in regional biodiversity dynamics. Despite the high level of turnover observed within these EE subunits, some level of ecologic stability occurs because abundant genera persist throughout entire EE subunits. Ordovician species in this study behaved relatively independently of other taxa and were not tightly integrated as suggested by the broadly overlapping taxon abundance curves, the shifting habitat preference of some taxa, the piecemeal turnover between EE subunits, and the continuous creation of new species associations due to background levels of turnover within EE subunits. Turnover within EE subunits was associated with relatively stable or only mildly fluctuating environments. Rapid turnover between EE subunits was caused by extreme perturbations to the regional or possibly global ocean-climate system.

Delayed biotic recovery from the end-Permian mass extinction has long been interpreted to result from environmental inhibition. Recently, evidence of more rapid recovery has begun to emerge, suggesting the role of environmental inhibition was previously overestimated. However, there have been few high-resolution taxonomic and ecological studies spanning the full Early and Middle Triassic recovery interval, leaving the precise pattern of recovery and underlying mechanisms poorly constrained. In this study, we document Early and Middle Triassic trends in taxonomic diversity, assemblage evenness, and size distribution of benthic foraminifers on an exceptionally exposed carbonate platform in south China. We observe gradual increases in all metrics through Early Triassic and earliest Middle Triassic time, with stable values reached early in the Anisian. There is little support in our data set for a substantial Early Triassic lag interval during the recovery of foraminifers or for a stepwise recovery pattern. The recovery pattern of foraminifers on the GBG corresponds well with available global data for this taxon and appears to parallel that of many benthic invertebrate clades. Early Triassic diversity increase in foraminifers was more gradual than in ammonoids and conodonts. However, foraminifers continued to increase in diversity, size, and evenness into Middle Triassic time, whereas diversity of ammonoids and conodonts declined. These contrasts suggest decoupling of recovery between benthic and pelagic environments; it is unclear whether these discrepancies reflect inherent contrasts in their evolutionary dynamics or the differential impact of Early Triassic ocean anoxia or associated environmental parameters on benthic ecosystems.

Extinction risk is inversely related to genus age (time since first appearance) in most intervals of the Phanerozoic marine fossil record, in apparent contradiction to the macroevolutionary Red Queen's Hypothesis, which posits that extinction risk is independent of taxon age. Age-dependent increases in the mean species richness and geographic range of genera have been invoked to reconcile this genus-level observation with the presumed prevalence of Red Queen dynamics at the species level. Here we test these explanations with data from the Paleobiology Database. Multiple logistic regression demonstrates that the association of extinction risk with genus age is not adequately explained by species richness or geographic range: there is a residual association between age and extinction risk even when range and richness effects are accounted for. Throughout most of the Phanerozoic the age selectivity gradient is highest among the youngest age cohorts, whereas there is no association between age and extinction risk among older age cohorts. Some of the apparent age selectivity of extinction in the global fauna is attributable to differences in extinction rate among taxonomic groups, but extinction risk declines with genus age even within most taxonomic orders. Notable exceptions to this pattern include the Cambrian-Ordovician, latest Permian, Triassic, and Paleocene intervals. The association of age with extinction risk could reflect sampling heterogeneity or taxonomic practice more than biological reality, but at present it is difficult to evaluate or correct for such biases. Alternatively, the pattern may reflect consistent extinction selectivity on some as-yet unidentified covariate of genus age. Although this latter explanation is not compatible with a Red Queen model if most genus extinctions have resulted from biological interactions, it may be applicable if most genus extinctions have instead been caused by recurrent physical disturbances that repeatedly impose similar selective pressures.

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.

The richness (number of species) and evenness (uniformity of species abundances) of death assemblages can differ from corresponding living communities due to processes such as between-habitat transport, environmental condensation, and differential taphonomic destruction. Analysis of 132 single-census live-dead comparisons of benthic molluscs from a variety of soft-bottom marine settings indicates that on average evenness does not differ greatly between live and dead assemblages, regardless of the particular depositional setting or grain size of associated sediment. However, individual death assemblages can deviate quite substantially from their corresponding living assemblages, especially if processed using a fine mesh. In addition, death assemblages collected using sieves with 2 mm mesh or coarser showed consistently and significantly greater evenness than corresponding living assemblages. These results are encouraging for broad-scale assessments of evenness in the fossil record based on the comparison of average values (rather than for individual assemblages) and where trends in evenness are the aim of the study.

Our live-dead comparisons of richness sample-size corrected by rarefaction revealed that death assemblages were on average ~1.45 times richer than the corresponding living assemblages regardless of rarefied size. In 63.6% of death assemblages both dead richness and dead evenness were greater than live, suggesting sufficient time-averaging to catch significant random or directional changes in the living community and/or introduction of individuals from outside the sampled habitat. In 12.9% of collections both dead richness and dead evenness were less than live, suggesting either rapid loss of dead shells so that dead diversity is depressed below the local living community or selective loss of taphonomically vulnerable taxa. In 18.2% of data sets dead richness was elevated but dead evenness was depressed relative to live: these are interpreted to reflect the addition of low-evenness allochthonous material. The remaining 4.5% of data sets had elevated dead evenness but depressed dead richness, suggesting that live and dead in this case may not be closely related.

In seven available time series, temporal volatility in living communities over 6–24 months was considerable but could not account for observed (mostly higher) evenness values in corresponding death assemblages, whose evenness and composition were quite stable in the few examined studies. A densely sampled spatial transect shows that changes in living-assemblage evenness along an environmental gradient were preserved in the corresponding death assemblages, although dead evenness at any location on the gradient was substantially higher than living evenness.

Many species of scleractinian reef corals deposit aragonite skeletons with cyclic bands of higher and lower density whose periodicity is annual. As these growth bands are frequently preserved in fossil corals, attention has been focused on their possible use as environmental indicators in as much as density variations may reflect changes in water temperature and/or light intensity. Relationships between skeletal growth banding and environmental parameters have been investigated by X-radiographic examination of 1488 specimens of modern reef corals representing 31 reef localities widely distributed over the Indo-Pacific and Caribbean regions. Average monthly seawater temperature and solar radiation data are available for each locality, and the date of collection is known for each sample. Of the 47 genera and subgenera included in the study, density variations were most pronounced in skeletons of Astreopora, Coscinarea, Cyphastrea, Favia (especially species with small corallites such as F. stelligera), Goniastrea, Hydnophora (massive forms only, e.g. M. microconos), Leptoria, Montastrea (especially M. annularis, with small corallites), Pavona, Polyastra, and Plesiastrea. Less useful are Diploastrea, Diploria, and Favites (which tend to have large corallite diameters), and Goniopora, Alveopora, Porites, Siderastrea, and Stephanaria (which frequently exhibit numerous secondary density variations within the annual cycle). Despite considerable variability among different individual corals from the same population, the average thickness of the skeletal growth bands is positively correlated with mean annual water temperature. By comparing characteristics of the outermost growth increment and the date of collection with monthly records of water temperature and solar radiation, it appears that maximum skeletal density is associated with those periods of the year when seawater temperature is above average.

Contours of whorl expansion rate (W), distance of the generating curve from the axis of coiling (D) and shape of the generating curve (S) were plotted from data on 988 Jurassic through Cretaceous ammonite species and 205 Jurassic through Tertiary nautilid species. The areas occupied by the ammonites and nautilids are largely nonoverlapping when plotted as WD and DS graphs.

The almost complete absence of overlap in Juro-Cretaceous ammonite and nautilid shell shape distributions is consistent with the view that ammonites and nautilids were dissimilar ecologically, and evolved different shell shapes in response to different adaptational needs. The terminal Cretaceous extinction of the ammonites may have opened up new opportunities for nautilid evolution during the Tertiary, because Tertiary nautilids are dominanted by moderately compressed, hydrodynamically efficient shell shapes which were rarely present among Jurassic and Cretacepus nautilids, but common among ammonites. The restriction of nautilids to a far more limited portion (compared to ammonites) of the theoretically available spectrum of planispiral shell shapes may be due in part to fabricational constraints imposed by the simple nautilid septum and septal suture. Nautilid shell shapes were limited to globular or slightly compressed to depressed shells, which are inherently strong due to pronounced shell wall curvature, and would be strengthened to only a small extent by buttressing from internal septa. Strongly depressed or compressed shell shapes, commonly evolved among ammonoids, had flattened shell regions which are inherently weak, and would be strengthened to a considerable extent by numerous thick, or strongly folded and fluted septa and septal sutures.

Biometric analyses are useful tools for the study of organisms, their phylogenetic affiliation, and the pattern and rate of their evolution. Various morphometric techniques have been developed to analyze morphological variation, but methodological choices are often made arbitrarily because quantitative comparisons are lacking or inconclusive. Here we address morphometric quantification of taxa with few unambiguously identifiable landmarks (<15), utilizing ornamented and unornamented gastropod shells. Support vector machines were applied to evaluate classification performances of landmark (LMA), elliptic Fourier (EFA), and semi-landmark analysis (SLM). This evaluation is based on the discrimination of between-group differences relative to within-group variation, and thus allows comparing how the techniques treat different types of biological information. The results suggest that EFA performs slightly better than SLM (and certainly LMA) in discerning a priori identified taxa with unornamented shells, but that SLM is significantly superior to other techniques for ornamented shells. Alignment and homology problems may cause the subtle variations in ornamentation to become blurred as noise in EFA, even though EFA is often cited to be able to deal with complex shapes. Performance of LMA depends entirely on how accurately the structure can be covered with landmarks. Guidelines in choosing a morphometric technique in diverse cases are provided.

The prosogyrous shape and flattened lunule of a typical clam shell form a blunt anterior, the function of which is related to the forward-and-back rocking motion of the shell in burrowing. Analysis of movies has revealed that each rocking motion of a morphologically typical clam, Mercenaria mercenaria (Linné), involves purely rotational movement, with no translational component. The clam is able to burrow by “walking” its way downward only because the axis of backward rotation lies to the anterior of the axis of forward rotation. Experiments with burrowing robots show that the blunt anterior serves to shift the axis of backward rotation anteriorly, thus aiding in downward progress. The prosogyrous condition and the rotational mechanism of burrowing are fundamental adaptations of burrowing clams and were apparently present in the ancestral bivalves of the Cambrian.

The Ordovician saw the transformation of marine benthic communities from the trilobite-based Cambrian Fauna to the brachiopod-dominated Paleozoic Fauna. An evaluation of the changing importance of trilobites during the Ordovician can be made from accurate assessments of taxonomic richness in various habitats. Here we present a new compilation of trilobite alpha diversity based on field collections and survey of the literature. The data indicate that trilobite species richness within nearshore, shallow subtidal, carbonate buildup and deep subtidal shelf environments was essentially constant between the Late Cambrian and the Late Ordovician. The alpha diversity patterns do not support the notion that trilobites became displaced from inner shelf environments during the Ordovician. Rather, the data are consistent with a decline in relative importance of the group through dilution as newly radiating invertebrate groups entered Ordovician paleocommunities. They also imply that direct interactions between elements of the Cambrian and Paleozoic faunas were not involved in the Ordovician reorganization of paleocommunities. Like many other major faunal transitions during the Phanerozoic, the Ordovician radiations appear to have been essentially non-competitive in nature.

Patterns of temporal variation of six characters in a Miocene stickleback (Gasterosteus doryssus) are presented. Most pairs of characters tend to be correlated, but these correlations account for only about 26% of the observed variation, and thus the characters are studied separately. All character state frequencies exhibit temporal heterogeneity, and their means have temporal trends. Regardless of these overall trends, reversals of the trends cause end members of four time series not to differ significantly from each other. We argue that most observed temporal variation represents intrapopulation evolutionary change. Although most time series have somewhat stepped patterns, complete stasis is absent. Gradual variation is predominant, and in one case is nearly linear, but very rapid evolutionary bursts are also seen. The most rapid stratigraphic change appears to represent a local extinction, followed by recolonization by a differentiated population, but another episode of rapid change probably represents evolution in situ by means of standard neo-Darwinian mechanisms, without involvement of “macromutations” or true saltation. The irregular patterns and great magnitude of phenotypic change that are observed indicate that conventional paleontological samples may miss important evolutionary phenomena and are not comparable to shorter-term evolution in extant populations.

The process of evolution hinders our ability to make large-scale ecological comparisons—such as those encompassing marine biotas spanning the Phanerozoic—because the compared entities are taxonomically and morphologically dissimilar. One solution is to focus instead on life habits, which are repeatedly discovered by taxa because of convergence. Such an approach is applied to a comparison of the ecological diversity of Paleozoic (Cambrian–Devonian) and modern marine biotas from deep-subtidal, soft-substrate habitats. Ecological diversity (richness and disparity) is operationalized by using a standardized ecospace framework that can be applied equally to extant and extinct organisms and is logically independent of taxonomy. Because individual states in the framework are chosen a priori and not customized for particular taxa, the framework fulfills the requirements of a universal theoretical ecospace. Unique ecological life habits can be recognized as each discrete, n-dimensional combination of character states in the framework. Although the basic unit of analysis remains the organism, the framework can be applied to other entities—species, clades, or multispecies assemblages—for the study of comparative paleoecology and ecology. Because the framework is quantifiable, it is amenable to analytical techniques used for morphological disparity. Using these methods, I demonstrate that the composite Paleozoic biota is approximately as rich in life habits as the sampled modern biota, but that the life habits in the modern biota are significantly more disparate than those in the Paleozoic; these results are robust to taphonomic standardization. Despite broadly similar distributions of life habits revealed by multivariate ordination, the modern biota is composed of life habits that are significantly enriched, among others, in mobility, infaunality, carnivory, and exploitation of other organisms (or structures) for occupation of microhabitats.

The modern structure of marine benthic ecosystems was largely established during the Jurassic and Early Cretaceous (200–100 Ma), a transition that has been termed the Mesozoic Marine Revolution (MMR). Although it has been suggested that the MMR marks an increase in the average energy consumption of marine animal ecosystems, this hypothesis has not been evaluated quantitatively. In this study, we integrate body size and abundance data from the fossil record with physiological data from living representatives to estimate mean per capita metabolic rates of tropical to subtropical assemblages of shallow-marine gastropods—a major component of marine ecosystems throughout the Meso-Cenozoic—both before and after the MMR. We find that mean per capita metabolic rate rose by ∼150% between the Late Triassic and Late Cretaceous and remained relatively stable thereafter. The most important factor governing the increase in metabolic rate was an increase in mean body size. In principle, this size increase could result from secular changes in sampling and taphonomic biases, but these biases are suggested to yield decreases rather than increases in mean size. Considering that post-MMR gastropod diversity is dominated by predators, the net primary production required to supply the energetic needs of the average individual increased by substantially more than 150%. These data support the hypothesis that benthic energy budgets increased during the MMR, possibly in response to rising primary productivity.

Methods currently used to estimate taxonomic extinction probabilities from fossil data generally assume that the probability of encountering a specimen in a particular stratum, given that the taxon was extant in the time period and location represented by the stratum, either equals 1.0 or else is a constant for all strata. Methods used to estimate taxonomic diversity (number of taxa) and speciation rate generally assume that encounter probabilities equal 1.0. We suspect that these assumptions are often false. Capture-recapture models were historically developed for estimation in the face of variable and unknown sampling probabilities. These models can thus be used to estimate parameters of interest from paleobiological data when encounter probabilities are unknown and variable over time. These models also permit estimation of sampling variances, and goodness-of-fit tests are available for assessing the fit of data to most models. Here we describe capture-recapture models which should be useful in paleobiological analyses and discuss the assumptions which underlie them. We illustrate these models with examples and discuss aspects of study design. We conclude that these models should prove useful in paleobiological analyses.

This paper presents a new means of interpreting the distribution of taxa among taxonomic lists. Traditionally, “similarity” indices have been used to compare lists, and “association” measures have been used to compare taxonomic distributions. It is argued that when sampling regimes are poorly understood, all similarity and association indices are unjustifiable. However, observations that taxa have overlapping (conjunct) or nonoverlapping (disjunct) distributions are universally meaningful. Because greater sampling can only increase the number of known conjunctions and because long lists serve as conjunctional Rosetta Stones, conjunction data sets can be far more reliable than the lists that generate them. One way to account for patterns of conjunction and disjunction is to create theoretical spaces composed of two distributional boundaries or edges for each taxon in each dimension. A parsimonious arrangement of edges implies distributions that always overlap when taxa are conjunct and infrequently overlap when taxa are disjunct. A procedure involving correspondence analysis is shown to minimize the number of implied disjunctions. The method is used to analyze a set of 271 Miocene large-mammal genus lists from the Great Plains region. Biostratigraphic and radiometric data demonstrate that the “best” arrangement of distributional edges is temporal and corresponds to an age-range chart. The edge sequence is calibrated to the radiometric time scale and used to compute a detailed genus-level diversity curve and to redefine the boundaries between Miocene Land-Mammal Ages and Subages in the Great Plains. Each of the Miocene Ages can be divided into two or three Subages. Diversity levels are apparently low throughout the Early Miocene and rise sharply during the Middle Miocene. Significant drops in diversity occurred at about 17.4, 11.6, 8.7, and 6.5 Ma. The base of the Barstovian corresponds to the base of the Middle Miocene, and the Clarendonian straddles the Middle-Late Miocene boundary.

Studies in the past 20 years have often interpreted size fluctuations in fossil mammal remains as a response to climatic change, in accord with Bergmann's rule. However, such paleoecological inference requires careful consideration of changes in community composition that could cause ecological character displacement or release. Recent size gradients of mammals should be screened for the possibility of character displacement if fossil size gradients are to be used as a “paleothermometer” for past climate. The use of teeth in paleontological studies as a measure of body size presents a further complication; for several carnivore guilds, it appears that competitive pressures act most strongly on tooth size. Teeth may therefore inaccurately estimate body size, even if body size accurately indicates climatic conditions. In the fossil record different species exhibit different size patterns under the same conditions of climatic change, and the same species may show diametrically opposite size fluctuations under similar conditions of climatic change, in different regions.

Using computer simulations and analytic calculations, we have evaluated whether conspicuous expansions and contractions of natural clades may have represented chance fluctuations that occurred while probabilities of speciation and extinction remained equal and constant. Our results differ from those of previous workers, who have not scaled generating parameters empirically at the species level. We have found that the waxing and waning of many real clades have almost certainly resulted from changes in probabilities of speciation and extinction. For some of these changes, likely explanations are evident. The emplacement of adaptive innovations, for example, has at times elevated probability of speciation. We conclude that chance factors have not played a dominant role in producing dramatic changes in standing diversity within speciose higher taxa.