The fossil record is incomplete in the sense that taxa for which there are no preserved finds are unrepresented. Because the probability that there is at least one preserved find is greater for long-lived taxa, failure to account for this effect will bias estimation of mean taxonomic duration. This paper describes maximum likelihood estimation of mean taxonomic duration, fossil preservation rate, and completeness of the local fossil record under a model in which duration has an exponential distribution and the locations of preserved finds follow a Poisson process. The estimates under this model have simple closed forms. It is also straightforward to construct confidence regions for the model parameters. The method is applied to data on 110 trilobite species from the Upper Cambrian—Lower Ordovician in Oklahoma.

Stratigraphic data are compared to morphologic data in terms of their fit to phylogenetic hypotheses for 29 data sets taken from the literature. Stratigraphic fit is measured using MacClade's stratigraphic character, which tracks the number of independent discrepancies between observed order and the order of occurrence that would be expected on the basis of a given phylogenetic hypothesis. Acceptance of a phylogenetic hypothesis despite such discrepancies requires ad hoc hypotheses concerning differential probabilities of preservation and recovery. These stratigraphic ad hoc hypotheses are treated as logically equivalent to morphologic ad hoc hypotheses of homoplasy. The retention index is used to compare the number of stratigraphic and morphologic ad hoc hypotheses required by given phylogenetic hypotheses. Each data set is subjected to five analyses, varying in the constraints imposed on the structure of the phylogenetic tree against which fit is measured. Analyses 1–4 compare the stratigraphic and morphologic retention indices using phylogenetic trees consistent with the morphologically most-parsimonious cladogram reported in the original study. Analysis 5 compares retention indices using the overall (stratigraphically and morphologically) most-parsimonious phylogenetic tree, which may be, but is not necessarily, consistent with the reported cladogram. Proceeding from Analysis 1 to Analysis 5, stratigraphic data are allowed greater influence in determining the structure of phylogenetic trees, with the trees in Analysis 1 derived without reference to the stratigraphic character and the trees in Analysis 5 derived from full interaction of stratigraphic and morphologic characters. Morphologic and stratigraphic retention indices for these 29 studies cannot be statistically distinguished in comparisons 3–5, suggesting very similar degrees of fit. The values of these retention indices are high, indicating a generally high level of congruence under these phylogenetic hypotheses. Significant gains (49%) in stratigraphic fit can be realized without significant loss (4%) in morphologic fit as the stratigraphic and morphologic evidence are both allowed to participate in constraining the structure of phylogenetic hypotheses. These results suggest that arguments based on alleged “noisiness” of stratigraphic data offer inadequate grounds for ignoring stratigraphic order in phylogenetic analysis. In terms of congruence, stratigraphic and morphologic data perform about equally well.

Computer models of the tail of Apatosaurus louisae show it could reach supersonic velocities, producing a noise analogous to the “crack” of a bullwhip. Similarity in tail structure suggests this was feasible for other diplodocids, and possibly for unrelated sauropods like Mamenchisaurus and the dicraeosaurids. Lengthening of caudal vertebrae centra between positions 18 and 25 is consistent with adaptation to the stresses generated by such tail motion, as is coossification of vertebrae via diffuse idiopathic skeletal hyperostosis (DISH), which occurs in the same region in about half the specimens. The noise produced may have been used for defense, communication, intraspecific rivalry, or courtship, in which case supersonic “cracking” may have been a sexually dimorphic feature. Comparisons with the club-bearing tails of the sauropods Shunosaurus lii and Omeisaurus tianfuensis show the diplodocid whiplash tail was not well adapted as a direct-impact weapon, bringing the tail-as-weapon hypothesis into doubt.

Paleontological completeness and stratigraphic ranges depend on extinction rate, origination rate, preservation rate, and the length of the interval of time over which observations can be made. I derive expressions for completeness and the distribution of durations and ranges as functions of these parameters, considering both continuous- and discrete-time models.

Previous work has shown that, if stratigraphic ranges can be followed indefinitely forward, and if extinction and preservation occur at stochastically constant rates, then extinction rate and preservability can be estimated from (1) discrete (binned) stratigraphic ranges even if data on occurrences within ranges are unknown, and (2) continuous ranges if the number of occurrences within each range is known. I show that, regardless of whether the window of observation is finite or infinite, extinction and preservation rates can also be estimated from (3) continuous ranges when the number of occurrences is not known, and (4) discrete ranges when the number of occurrences is not known. One previous estimation method for binned data involves a sample-size bias. This is circumvented by using maximum likelihood parameter estimation. It is worth exploiting data on occurrences within ranges when these are available, since they allow preservation rate to be estimated with less variance. The various methods yield comparable parameter estimates when applied to Cambro-Ordovician trilobite species and Cenozoic mammal species.

Stratigraphic gaps and variable preservation affect stratigraphic ranges predictably. In many cases, accurate parameter estimation is possible even in the face of these complications. The distribution of stratigraphic ranges can be used to estimate the sizes of gaps if their existence is known.

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.

A generalized method for calculating confidence intervals on the position of the true end point of a stratigraphic range when the distributions of fossil horizons is nonrandom is presented. The method requires a quantitative measure of collecting and/or preservation biases with stratigraphic position. This fossil recovery potential function may be based on (among other variables) bedding-plane surface areas, or, given a water depth curve, an a priori estimate of the preservation potential with water depth. The approach assumes that the observed distribution of fossil horizons is consistent with the distribution predicted by the fossil recovery potential function, an assumption that must be tested before the method is applied. Unlike previous methods for calculating confidence intervals on the end points of stratigraphic ranges, this method may be applied when the number of fossil horizons is correlated with stratigraphic position. The approach should only be applied to sections that have been sampled continuously, or approximately continuously. Its efficacy will depend on how accurately fossil recovery potentials can be determined. A method is also presented for estimating the probability that a species became extinct during a major hiatus in the rock record.

Use of quantitative statistical tests can show that there is generally good congruence between estimated cladistic hypotheses of relationship and observed stratigraphy. A data set of 376 cladograms of fishes, continental tetrapods, and echinoderms was tested using three metrics, Spearman Rank Correlation (SRC), Relative Completeness Index (RCI), and Stratigraphic Consistency Index (SCI), to explore the relationships between the indices and differences in results among the three groups of organisms.

There is a strong relationship between SCI and SRC, since both tests measure the same aspect of the fossil record. There is no relationship between RCI and either SCI or SRC. There is a highly significant relationship, as expected, between SRC coefficients and the number of taxa in a cladogram, but no such relationship for RCI or SCI (except in fishes). There is no significant relationship between any of the indices and either the number of internal nodes or tree balance.

Echinoderms show the best stratigraphic consistency of nodes, while continental tetrapods have the best matching of stratigraphic age and cladistic node order. Fishes have the worst match of age and clade ranks, but they do have the most complete fossil record as measured by the RCI. They are followed by echinoderms, and then continental tetrapods, which have the least complete record. This seems to show that life in an aquatic environment leads, in general, to a more complete fossil record.

The ammonoid order Prolecanitida constitutes a relatively small (43 genera, ~250 species) but long-ranging lineage (Lower Carboniferous—Triassic, ~108 m.y.), which narrowly survived the P/Tr extinctions and provided the stock from which were derived all later Mesozoic ammonoids. Prolecanitids were a minority among Late Paleozoic ammonoids, which were dominated by the Goniatitida, and showed many features that set them far apart from their contemporaries, including (1) long-term, gradual changes in shell geometry (W-D-S); (2) the most strongly constrained morphospace of any Paleozoic ammonids examined to date; (3) an eight-fold increase in mean suture complexity (three times that of Pennsylvanian goniatitids); (4) high correlations between shell geometry, shell and septal thickness, and suture complexity; (5) short body chambers and, as a consequence, high aperture orientations; (6) indications that cameral liquid may have been used for buoyancy control; and (7) a genus longevity that averaged 14.7 m.y. compared with 5.7 m.y. in Upper Carboniferous goniatitids, and that appears to have been unrelated to suture complexity. Prolecanitids showed a pervasive tendency to increase suture complexity (in the clade as a whole as well as within subclades and in more than 90 percent of ancestor-descendant genera), thus arguing a case for a driven complexity trend. The uniqueness of the prolecanitids calls into question whether they and their Mesozoic descendants, ceratites and ammonites, were strictly analogous to Paleozoic goniatites.

The stratigraphic distribution of fossil species contains potential information about phylogeny because some phylogenetic trees are more consistent with the distribution of fossils in the rock record than others. A maximum likelihood estimator of phylogeny is derived using an explicit mathematical model of fossil preservation. The method assumes that fossil preservations within lineages follow an independent Poisson process, but can be extended to include other preservation models. The performance of the method was examined using Monte Carlo simulation. The performance of the maximum likelihood estimator of topology increases with an increase in the preservation rate. The method is biased, like other methods of phylogeny estimation, when the rate of fossil preservation is low; estimated trees tend to be more asymmetrical than the true tree. The method appears to perform well as a tree rooting criterion even when preservation rates are low. We suggest several possible extensions of the method to address other questions about the nature of fossil preservation and the process of speciation and extinction over time and space.

Two living forms of Globigerinella siphonifera (d'Orbigny), presently identified as Type I and Type II, can easily be distinguished and collected by SCUBA divers because of differences in appearance, arrangement of the rhizopodial network, and the presence or absence of commensals. Additional biological differences are apparent from laboratory culture experiments; Type I individuals survive significantly longer than Type II under conditions of darkness and starvation and have significantly slower chamber formation rates. Stable isotopic analyses of Types I and II also reveal notable differences, with Type I consistently yielding more negative δ18O and δ13C values. Results of Mg/Ca ratio analyses indicate that Type II specimens precipitated their shells in slightly cooler (deeper) surface waters than Type I specimens. These observations and results from DNA sequencing unequivocally demonstrate that G. siphonifera Types I and II should be regarded as biological sister species.

Contrarily, biometric analysis of the empty shells reveals few significant differences between G. siphonifera Types I and II. Of all the features measured from X-ray and SEM images of serially dissected specimens, only shell porosity yields readily discernible differences, with Type I adult chambers averaging 10–20% porosity and Type II adult chambers averaging 4–7% porosity. Statistically significant differences between Type I and II populations are revealed in maximum test diameter (Type I is typically larger) and coiling (Type I is typically more evolute), but these differences do not justify species level distinction of Types I and II using traditional paleontological species concepts.

On the basis of the above evidence, and since all specimens were collected at the same location at ∼3–8 m water depth, we conclude that G. siphonifera Types I and II are living examples of cryptic speciation, whereby biological speciation has occurred in the absence of discernable change in shell morphology. However, it is not clear when or where this speciation took place. Preliminary study of deep-sea cores from the Caribbean and Pacific sides of the Isthmus of Panama reveals a predominance of specimens with Type II porosity values, with rare occurrence of specimens yielding Type I porosity values. Systematic downcore measurement of shell porosity and tightness of coiling needs to be extended back to the middle Miocene, when G. siphonifera first appeared, to determine the timing of the Type I and II morphological divergence.

Postulated mechanisms for reproductive isolation and speciation of Types I and II include alloparapatric, depth parapatric, and sympatric speciation. These models could be tested if further analysis of fossil G. siphonifera shells allows determination of the timing of speciation, the preferred depth distribution, and the history of geographic distribution of Types I and II.

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.

The extreme morphological simplicity of lingulide brachiopod shells makes them particularly useful for investigating the species-level taxonomic resolution of the fossil record as well as the relationships between taxonomy, morphological complexity, and evolutionary rates. Lingulides have undergone little change in shell morphology and have had low taxonomic diversity since the Paleozoic. Is this pattern an evolutionary phenomenon or an artifact of the shell's simplicity? Multivariate methods were used to establish morphogroups among seven populations of four extant species of Glottidia. Six characters (three shell dimensions and three internal septa) were measured for 162 specimens from field and museum collections. All populations follow similar allometric trajectories: internal septa display positive allometry and shell dimensions display negative allometry. The allometric pattern may reflect D'Arcy Thompson's Principle of Similitude. Principal component analysis does not reveal any distinct clusters in Glottidia morphospace but suggests that some differences independent from ontogeny exist among the populations. Size-free canonical variate analysis indicates the presence of five size-invariant groups that are statistically distinct. Bootstrap-corrected error rates indicate that four specimens are enough to classify a sample correctly at α = 0.05 and eight specimens at α = 0.01. The groups are consistent with neontological classification with the exception of two populations of G. pyramidata identified by discriminant analysis as two distinct groups. The size-free morphogroups reflect geographic separation rather than ontogenetic or substrate differences among the populations.

Despite the morphological simplicity of the shell, size-free multivariate analysis of Glottidia delineates groups that offer taxonomic resolution comparable with the neontological classification. The method offers a promising tool for identifying natural morphogroups on the basis of few morphological characters. Moreover, the agreement between neontological taxonomy and the morphogroups suggests that the size-free approach can be applicable for evaluating the reality of the low diversity and turnover rates observed in the fossil record of lingulide brachiopods (= Family Lingulidae). Assuming that the neontological species of Glottidia are biologically meaningful, this study shows that morphological simplicity of lingulides does not necessarily result in taxonomic underresolution. Our analysis, as well as several previous case studies, suggests that taxonomic diversity and turnover rates do not have to be dependent on the morphological complexity of preservable parts. In many cases, when rigorous quantitative methods are employed, the differences in the rates of morphological evolution may be a real evolutionary phenomenon and not artifacts of morphological complexity.

Morphological analysis of four higher taxa of fossil marine invertebrates shows that, over the history of paleontology, there is no general tendency for morphologically extreme or modal species and genera to be described preferentially early or late. Reconstructing the expected evolutionary sequences of morphological disparity that would have been estimated at various times during the past century and a half reveals features that are sensitive to sampling (for example, peak trilobite disparity in the Ordovician, peak of post-Paleozoic crinoid disparity in the Triassic, and peak blastoid disparity in the Permian), as well as more robust features (for example, increase in trilobite disparity from the Cambrian to the Ordovician, continued increase in trilobite disparity despite a drop in taxonomic diversity after the Early Ordovician, decrease in blastoid disparity from the Devonian to the Carboniferous, and increase in crinoid disparity from the Jurassic to the Cretaceous followed by decline during the Cretaceous). Although we still have much to learn about the evolution of form, in many respects our view of the history of biological diversity is mature.

Disturbances have drastically altered Caribbean coral reefs over the past two decades. Acropora cervicornis (staghorn coral), which predominated at intermediate depths (5—25 m) from the 1950s through the 1970s, has virtually disappeared from most reef environments. Other coral species have declined as well, and the cover of macroalgae has increased. In apparent contrast, fossil reef sequences suggest that the species composition and zonation of coral assemblages did not change during the Pleistocene and Holocene. One interpretation of these observations is that coral species persisted on Caribbean reefs for hundreds of thousands of years as components of tightly integrated communities, and that a rare or unique combination of disturbances led to the synchronous decline of A. cervicornis and other corals throughout the region. The hypotheses of (1) community integration and (2) a unique, recent community transition, were tested by ecological and paleoecological observations in the shelf lagoon of the Belizean Barrier Reef.

The reef growing along the flanks of Channel Cay, a lagoonal shoal, was monitored by point counts along transects over a ten-year period (1986—95). This reef was covered primarily by A. cervicornis at 3—15 m depth until the late 1980s. After 1986, A. cervicornis experienced a mass mortality from White Band Disease, an epizootic of presumed bacterial origin. The cover of A. cervicornis dropped from ~70% in 1986 to nearly 0% in 1993. Agaricia spp. (lettuce corals) responded opportunistically to the availability of free space in the form of A. cervicornis skeletal rubble. Agaricia, which had been a minor constituent of the sessile biota (10% cover in 1986), replaced A. cervicornis as the most common occupant of space on the reef (56% cover in 1995). The percent cover of other coral species and macroalgae remained low throughout the ten-year period. Similar changes were observed on other reefs over an area of at least 250 km2.

The Acropora-to-Agaricia transition left a clear signature in the sedimentary record. Trenches dug into the reef at Channel Cay revealed the accretion of a layer of Agaricia rubble with a mean thickness of 22 cm in the decade after 1986. Due to the unconsolidated, uncompacted nature of the reef sediments, evidence of previous Acropora-to-Agaricia transitions should have been visible in the fossil record as vertical accumulations of A. cervicornis branches interrupted by layers of imbricated Agaricia rubble. Coring studies at Channel Cay revealed that no other Agaricia layers were deposited during at least the past 3800 years; the recent transition was unique on a time scale of millennia. This result supports the contention that excursions from the Acropora-dominated situation are unusual in the history of Channel Cay and nearby reefs. However, the dynamics of the transition do not support the community integration hypothesis for the Channel Cay reef, indicating instead that different coral taxa in this assemblage responded differently, or not at all, to a large-scale biotic disturbance. The community transition also underscores the potential for biological factors in general, and disease in particular, to alter the composition of ecological communities and their sedimentary remains.

Computer-generated searches through hypothetical fitness landscapes for progressively more fit variants were used to assess the effects of developmental barriers (mimicked by barring specific types of morphological transformations) and abruptly shifting environmental conditions (mimicked by sudden shifts in how fitness was defined) on the number and accessibility of optimal phenotypes. Relative fitness was defined in terms of maximizing light interception, mechanical stability, or reproductive success, or minimizing surface area, or optimizing the performance of various combinations of these tasks. Developmentally obstructed and unobstructed walks located, on average, equivalent numbers of phenotypic optima. The number of optima identified by both kinds of walks increased in proportion to the number of simultaneously performed tasks used to measure fitness. Walks passing from more complex to less complex fitness landscapes located more optima than walks passing through unchanging, stable landscapes. The model thus suggests that there are no a priori reasons to assume that (1) the morphological options available to adaptive evolution become more restrictive as biological complexity increases, (2) “developmental barriers” necessarily restrain a lineage from evolving well-adapted morphologies, and (3) generalist organisms are less successful than specialists. Also, because the number and accessibility of fitness peaks were proportional to the complexity of fitness landscapes, the model predicts that the probability of cladogenesis will increase as landscape complexity increases, while anagenesis will be encouraged when fitness is defined in terms of performing one or a few tasks simultaneously.

The evolution of septal complexity in fossil ammonoids has been widely regarded as an adaptive response to mechanical stresses imposed on the shell by hydrostatic pressure. Thus, septal (and hence sutural) complexity has been used as a proxy for depth: for a given amount of septal material greater complexity permitted greater habitat depth. We show that the ultimate septum is the weakest part of the chambered shell. Additionally, finite element stress analyses of a variety of septal geometries exposed to pressure stresses show that any departure from a hemispherical shape actually yields higher, not lower, stresses in the septal surface. Further analyses show, however, that an increase in complexity is consistent with selective pressures of predation and buoyancy control. Regardless of the mechanisms that drove the evolution of septal complexity, our results clearly reject the assertion that complexly sutured ammonoids were able to inhabit deeper water than did ammonoids with simpler septa. We suggest that while more complexly sutured ammonoids were limited to shallower habitats, the accompanying more complex septal topograhies enhanced buoyancy regulation (chamber emptying and refilling), through increased surface tension effects.

The exceedingly rich middle Pleistocene mammalian fauna from the classic Ensenadan Tarija basin in southern Bolivia contains a diversity of medium to large-bodied herbivores consisting of both endemic (†Toxodontia, †Litopterna, Xenarthra) and immigrant (Rodentia, Proboscidea, Perissodactyla, and Artiodactyla) taxa. In order to characterize feeding ecology and niche differences, a suite of morphological characters was measured for each of 13 species of herbivorous mammals from the Pleistocene of Tarija; these were combined with carbon isotopic results from tooth enamel. (The Xenarthra were excluded from this study because they lack tooth enamel.)

Several different bivariate and multivariate combinations of characters can be used to characterize the feeding adaptations, niches, and guild composition of the Tarija mammalian herbivores. During the Pleistocene the browsing guild in the Tarija basin is interpreted to include the tapir (Tapirus tarijensis), extinct llama (Palaeolama weddelli), peccary (Tayassu sp.), and deer (Hippocamelus sp.). The mixed-feeding guild included two horse species (Hippidion principale and Onohippidium devillei), litoptern (Macrauchenia patachonica), and capybara (Neochoerus tarijensis). The grazing guild included the numerically dominant horse (Equus insulatus), two lamine species (Lama angustimaxilla and cf. Vicugna, provicugna), notoungulate (Toxodon platensis), and gomphothere proboscidean (Cuvieronius hyodon). The grazing guild has the widest range of body sizes relative to the two other guilds. Closely related sympatric species within the Equidae and Camelidae differentiate their niches from one another using a combination of body size, feeding ecology, and probably local habitat. Most of the paleoecological reconstructions resulting from this combined morphological and isotopic analysis corroborate previous studies based primarily on morphology; there are, however, some notable surprises.

Late Eocene foraminiferal extinction shows diverse patterns of selective morphologic and latitudinal extinction. Taxa with discoidal shape, calcareous tests, and narrow and low-latitudinal ranges are at significantly greater risk of extinction. Elevated extinction intensities in calcareous tests are mainly due to the presence of larger benthic foraminifera that evolved in late Paleocene and diversified through the lower to middle Eocene. Selectivity of late Eocene foraminiferal extinction indicates that this extinction event was not a globally uniform event. Although this result does not verify an extraterrestrial impact or any other proposed cause of extinction, it does constrain the causes of late Eocene extinction. Furthermore, the geography of late Eocene foraminiferal extinction, and previously studied Cenomanian/Turonian extinction, demonstrates that mass extinctions exhibit different patterns of selectivity.

We used radiocarbon ages on dead Holocene shells of the venerid bivalve Chione spp. to investigate how time-averaging and taphonomy in shallow marine benthic assemblages vary with sedimentary and tectonic setting. We compared shells collected from the sediment surface in five depositional environments from two regions of the Gulf of California, Mexico: Bahía Concepción, a young faulted rift basin with high rates of terrigenous and carbonate sedimentation; and Bahía la Choya, an intertidal system along a sediment-starved shelf. Frequency distributions of shell ages in all environments form a hollow curve, with a mode at young ages and a long tail toward older ages. This pattern suggests that shells are added to the taphonomically active zone (TAZ) at roughly constant rates (via continuous shell deaths), and removed from the TAZ at random, either through destruction or by achieving final burial. Shell half-lives (the amount of time to remove half the shells from the TAZ) provide a comparative measure of time-averaging. Time-averaging varies with sedimentary and tectonic setting. The lowest amounts of time-averaging (shell half-lives of 90 to 165 years) occur in Bahía Concepción, where rapid rates of terrigenous sedimentation (on fan-deltas) and carbonate sedimentation (in pocket bays) bury shells rapidly. Time-averaging is higher in the sediment-starved environments of Bahía la Choya (shell half-lives of 285 to 550 years). The highest amounts of time-averaging occur the inner tidal flats of Bahía la Choya (shell half-life of 550 years). Here the conjunction of low sedimentation rates with low rates of shell destruction (due to periodic tidal emergence) permits shells to persist in the TAZ for very long time spans.

There is no systematic relationship between a shell's age and its taphonomic condition (taphonomic grade) in any environment, probably because of the complex and random nature of burial-exhumation in the TAZ. Age variance tends to increase with increasing taphonomic alteration: highly altered shells range in age from young to several thousand years old, while less altered shells are mostly young. The correspondence between time-averaging and the taphonomic condition of entire shell assemblages is also weak, but might be resolved with further study.

These results provide quantitative data on time-averaging in benthic assemblages as a function of sedimentary and tectonic setting, and suggest some guidelines for facies appropriate for particular studies. Shallow marine rift basins like Bahía Concepción can potentially contain within-horizon fossil assemblages representing time spans of only a few hundred years—time resolution often beyond reach in paleontology. In contrast, sediment-starved shelf habitats like Bahía la Choya are unlikely to yield assemblages with time resolution finer than several thousands of years.