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.

The late Neogene was a time of major environmental change in Tropical America. Global cooling and associated oceanographic reorganization and the onset and intensification of glaciation in the Northern Hemisphere during the past ten million years coincided with the uplift of the Central American isthmus and resulting changes in regional oceanographic conditions. Previous analyses of patterns of taxonomic turnover and the shifting abundances of major ecological guilds indicated that the regional shallow-water marine biota responded to these environmental changes through extinction and via a restructuring of local benthic food webs, but it is not clear whether this ecological response had an effect on the diversity of molluscan assemblages in the region. Changes in regional and local diversity are often used as proxies for similar ecological response to environmental change in large-scale paleontological studies, but a clear relationship between diversity and ecological function has rarely been demonstrated in marine systems dominated by mollusks. To explore this relationship, we have compiled a data set of the stratigraphic and environmental distribution of genera of mollusks in large new collections of fossil specimens from the late Neogene and Recent of the southwestern Caribbean. Analysis of a selection of ecological diversity measures indicates that within shelf depths, assemblages from deeper water (51–200 m) were more diverse than shallow-water (<50 m) assemblages in the Pliocene. Lower diversity for shallow-water assemblages is caused by increased dominance of a few superabundant taxa in each assemblage. This implies that studies of diversity of shelf benthos need to control for relatively fine scaled environmental conditions if they are to avoid interpreting artifacts of uneven sampling as true change of diversity. For shallow-water assemblages only, there was significant increase in local and regional diversity of bivalve assemblages after the late Pliocene. No parallel increase in gastropods could be detected, but this likely is because sample size was inadequate for documenting the diversity of gastropod assemblages following a steep post-Pliocene decline of average gastropod abundance. Both the increasing bivalve diversity and the decrease in average abundance of gastropod taxa correspond to an interval of increasing carbonate deposition and reef building in the region, and are likely a result of increased fine-scale habitat heterogeneity controlled by the local distribution of carbonate buildups. Each of these results demonstrates that documenting the ecological response of tropical marine ecosystems to regional environmental change requires a large volume of fine-scaled samples with detailed paleoenvironmental control. Such data sets are rarely available from the fossil record.

Cheek teeth of some mammalian herbivores exhibit pronounced changes in occlusal size and shape through wear, purportedly caused by strong curvature. Such changes are extreme in the upper cheek teeth of extinct, dentally archaic lagomorphs. Morphologic and taxonomic turnover in lagomorphs suggests that these dentally archaic forms may have been unable to develop hypselodont (ever-growing) cheek teeth. This study investigates how the interaction of tooth shape and wear can cause occlusal size and shape changes, and potentially impose structural constraints on crown height. These constraints may help explain extinction of mammals with teeth like archaic lagomorphs, evolution and diversification of other mammalian herbivores during the late Miocene, and the relative paucity of hypsodont cheek tooth shapes in extant mammals.

I first quantify two-dimensional curvature accounting for shape differences observed in hypsodont teeth, P4s of the archaic lagomorphs Russellagus and Hesperolagomys, which exhibit pronounced change with wear, and Ondatra lower incisors, which show minimal change with wear. Using this quantification, I generate theoretical curvature morphologies and describe a geometric model of tooth wear that generates values for qualitative and quantitative aspects of the occlusal surface at different wear stages. Modeled results of wear surface topography and dimensions closely correspond to observed patterns in Russellagus, Hesperolagomys, and Ondatra. Model results on wear in theoretical tooth morphologies identify two major shape factors influencing wear: orientation of the wear surface (incisor-like or cheek-tooth-like), and tooth curvature (“concentric” or “nonconcentric”). Modeled wear also suggests two geometric constraints on crown height. Teeth with nonconcentric curvatures can have crown height limited by potential tooth area. “Incomplete wear” in any tooth can present severe constraints on increasing crown height, causing structurally untenable morphologies in very tall-crowned to hypselodont teeth.

We present a new three-dimensional theoretical ecospace for the ecological classification of marine animals based on vertical tiering, motility level, and feeding mechanism. In this context, analyses of a database of level-bottom fossil assemblages with abundance counts demonstrate fundamental changes in marine animal ecosystems between the mid-Paleozoic (461–359 Ma) and late Cenozoic (23–0.01 Ma). The average local relative abundance of infaunal burrowers, facultatively motile animals, and predators increased, whereas surface dwellers and completely non-motile animals decreased in abundance. Considering tiering, motility, and feeding together, more modes of life had high to moderate average relative abundance in the Cenozoic than in the Paleozoic. These results are robust to the biasing effects of aragonite dissolution in Paleozoic sediments and to heterogeneities in the latitudinal and environmental distributions of collections. Theoretical ecospace provides a unified system for future analyses of the utilization of ecologic opportunities by marine metazoa.

This paper takes an alternative approach to the problem of inferring patterns of phenotypic evolution in the fossil record. Reconstructing temporal biological signal from noisy stratophenetic data is an inverse problem analogous to subsurface reconstructions in geophysics, and similar methods apply. To increase the information content of stratophenetic series, available geological data on sample ages and environments are included as prior knowledge, and all inferences are conditioned on the uncertainty in these geological variables. This uncertainty, as well as data error and the stochasticity of fossil preservation and evolution, prevents any unique solution to the stratophenetic inverse problem. Instead, the solution is defined as a distribution of model parameter values that explain the data to varying degrees. This distribution is obtained by direct Monte Carlo sampling of the parameter space, and evaluated with Bayesian integrals. The Bayesian inversion is illustrated with Miocene stratigraphic data from the ODP Leg 174AX Bethany Beach borehole. A sample of the benthic foraminifer Pseudononion pizarrensis is used to obtain a phenotypic covariance matrix for outline shape, which constrains a model of multivariate shape evolution. The forward model combines this evolutionary model and stochastic models of fossil occurrence with the empirical sedimentary record to generate predicted stratophenetic series. A synthetic data set is inverted, using the Neighbourhood Algorithm to sample the parameter space and characterize the posterior probability distribution. Despite small sample sizes and noisy shape data, most of the generating parameter values are well resolved, and the underlying pattern of phenotypic evolution can be reconstructed, with quantitative measures of uncertainty. Inversion of a stratigraphic series into a time series can significantly improve our perception and interpretation of an evolutionary pattern.

Paleobiological diversity is often expressed as α (within-sample), β (among-sample), and γ (total) diversities. However, when studying the effects of extinction on diversity patterns, only variations in α and γ diversities are typically addressed. A null model that examines changes in β diversity as a function of percent extinction is presented here.

The model examines diversity in the context of a hierarchical sampling strategy that allows for the additive partitioning of γ diversity into mean α and β diversities at varying scales. Here, the sampling hierarchy has four levels: samples, beds, facies, and region; thus, there are four levels of α diversity (α₁, α₂, α₃, α₄) and three levels of β diversity (β₁, β₂, and β₃). Taxa are randomly assigned to samples within the hierarchy according to probability of occurrence, and initial mean α and β values are calculated. A regional extinction is imposed, and the hierarchy is resampled from the remaining extant taxa. Post-extinction mean α and β values are then calculated.

Both non-selective and selective extinctions with respect to taxon abundance yield decreases in α, β, and γ diversities. Non-selective extinction with respect to taxon abundance shows little effect on diversity partitioning except at the highest extinction magnitudes (above 75% extinction), where the contribution of α₁ to total γ increases at the expense of β₃, with β₁ and β₂ varying little with increasing extinction magnitude. The pre-extinction contribution of α₁ to total diversity increases with increased probabilities of taxon occurrence and the number of shared taxa between facies. Both β₁ and β₂ contribute equally to total diversity at low occurrence probabilities, but β₂ is negligible at high probabilities, because individual samples preserve all the taxonomic variation present within a facies. Selective extinction with respect to rare taxa indicates a constant increase in α₁ and constant decrease in β₃ with increasing extinction magnitudes, whereas selective extinction with respect to abundant taxa yields the opposite pattern of an initial decrease in α₁ and increase in β₃. Both β₁ and β₂ remain constant with increasing extinction for both cases of selectivity. By comparing diversity partitioning before and after an extinction event, it may be possible to determine whether the extinction was selective with respect to taxon abundances, and if so, whether that selectivity was against rare or abundant taxa.

Field data were collected across a Late Ordovician regional extinction in the Nashville Dome of Tennessee, with sampling hierarchy similar to that of the model. These data agree with the abundant-selective model, showing declines in α, β, and γ diversities, and a decrease in α₁ and increase in β₃, which suggests this extinction may have targeted abundant taxa.

Tectonic deformation is an important part of the taphonomic histories of many fossils. Although the effects of deformation, and methods to remove those effects, have been a subject of inquiry for over a century, systematic testing under known parameters has never been used to determine how the effects of deformation and the performance of retrodeformation techniques might vary. Comparative studies of morphology depend on the accurate estimation of variance-covariance structure, so an understanding of the effects of retrodeformation on covariance structure is important in assessing the utility of these methods. Here we address these issues by using geometric morphometric simulations. Nondeformed data sets were generated from specimens of the extant turtle Emys marmorata, which were known by definition to be nondeformed, and which possess a known ontogenetic signal. Deformation was simulated by applying a combination of uniform shear and uniform compression/dilation to the data. Data were retrodeformed by reflection and averaging of bilaterally symmetric landmarks, use of a principal components analysis to identify a deformation component of shape variation, and removal of the affine component of shape variation among specimens. Deformation increased the amount of variance in the data, as well as altering the variance structure. However, low to moderate levels of deformation did not prevent the confident recovery of the known ontogenetic signal in some cases. The tested retrodeformation techniques did not work well. They either removed too little or too much variance from the data, and provided little improvement in variance structure. Retrodeformation often did not improve our ability to extract the ontogenetic signal from the data, and in some cases introduced an artifactual relationship between size and shape. All of the scrutinized methods showed some properties, such as reducing variance or producing visually appealing images of specimens, that could make them appear to be working in cases where the correct biological signal is not known. This emphasizes the need for simulation testing in the development and evaluation of retrodeformation techniques.

Echinoderms have long been characterized by the presence of ambulacra that exhibit pentaradiate symmetry and define five primary body axes. In reality, truly pentaradial ambulacral symmetry is a condition derived only once in the evolutionary history of echinoderms and is restricted to eleutherozoans, the clade that contains most living echinoderm species. In contrast, early echinoderms have a bilaterally symmetrical 2–1–2 arrangement, with three ambulacra radiating from the mouth. Branching of the two side ambulacra during ontogeny produces the five adult rays. During the Cambrian Explosion and Ordovician Radiation, some 30 clades of echinoderms evolved, many of which have aberrant ambulacral systems with one to four rays. Unfortunately, no underlying model has emerged that explains ambulacral homologies among disparate forms. Here we show that most Paleozoic echinoderms are characterized by uniquely identifiable ambulacra that develop in three distinct postlarval stages. Nearly all “aberrant” echinoderm morphologies can be explained by the paedomorphic ambulacra reduction (PAR) model through the loss of some combination of these growth stages during ontogeny. Superficially similar patterns of ambulacral reduction in distantly related clades have resulted from the parallel loss of homologous ambulacra during ontogeny. Pseudo-fivefold symmetry seen in Blastoidea and the true fivefold symmetry seen in Eleutherozoa result from great reduction and total loss, respectively, of the 2– 1–2 symmetry early in ontogeny. These ambulacral variations suggest that both developmental and ecological constraints affect the evolution of novel echinoderm body plans.