Understanding what drives global diversity requires knowledge of the processes that control diversity and turnover at a variety of geographic and temporal scales. This is of particular importance in the study of mass extinctions, which have disproportionate effects on the global ecosystem and have been shown to vary geographically in extinction magnitude and rate of recovery.

Here, we analyze regional diversity and turnover patterns for the paleocontinents of Laurentia, Baltica, and Avalonia spanning the Late Ordovician mass extinction and Early Silurian recovery. Using a database of genus occurrences for inarticulate and articulate brachiopods, bivalves, anthozoans, and trilobites, we show that sampling-standardized diversity trends differ for the three regions. Diversity rebounded to pre-extinction levels within 5 Myr in the paleocontinent of Laurentia, compared with 15 Myr or longer for Baltica and Avalonia. This increased rate of recovery in Laurentia was due to both lower Late Ordovician extinction rates and higher Early Silurian origination rates relative to the other continents. Using brachiopod data, we dissected the Rhuddanian recovery into genus origination and invasion. This analysis revealed that standing diversity in the Rhuddanian consisted of a higher proportion of invading taxa in Laurentia than in either Baltica or Avalonia. Removing invading genera from diversity counts caused Rhuddanian diversity to fall in Laurentia. However, Laurentian diversity still rebounded to pre-extinction levels within 10 Myr of the extinction event, indicating that genus origination rates were also higher in Laurentia than in either Baltica or Avalonia. Though brachiopod diversity in Laurentia was lower than in the higher-latitude continents prior to the extinction, increased immigration and genus origination rates made it the most diverse continent following the extinction. Higher rates of origination in Laurentia may be explained by its large size, paleogeographic location, and vast epicontinental seas. It is possible that the tropical position of Laurentia buffered it somewhat from the intense climatic fluctuations associated with the extinction event, reducing extinction intensities and allowing for a more rapid rebound in this region. Hypotheses explaining the increased levels of invasion into Laurentia remain largely untested and require further scrutiny. Nevertheless, the Late Ordovician mass extinction joins the Late Permian and end-Cretaceous as global extinction events displaying an underlying spatial complexity.

Previous analyses of the history of Phanerozoic marine biodiversity suggested that the post-Paleozoic increase observed at the family level and below was caused, in part, by an increase in global provinciality associated with the breakup of Pangea. Efforts to characterize the Phanerozoic history of provinciality, however, have been compromised by interval-to-interval variations in the methods and standards used by researchers to calibrate the number of provinces. With the development of comprehensive, occurrence-based data repositories such as the Paleobiology Database (PaleoDB), it is now possible to analyze directly the degree of global compositional disparity as a function of geographic distance (geo-disparity) and changes thereof throughout the history of marine animal life. Here, we present a protocol for assessing the Phanerozoic history of geo-disparity, and we apply it to stratigraphic bins arrayed throughout the Phanerozoic for which data were accessed from the PaleoDB. Our analyses provide no indication of a secular Phanerozoic increase in geo-disparity. Furthermore, fundamental characteristics of geo-disparity may have changed from era to era in concert with changes to marine venues, although these patterns will require further scrutiny in future investigations.

With species found throughout both marine and fresh waters, the diatom order Thalassiosirales is one of the most phylogenetically and ecologically diverse lineages of planktonic diatoms. A clear understanding of the timescale of Thalassiosirales evolution would provide novel insights into the rates and patterns of species diversification associated with major habitat shifts, as well as provide valuable context for understanding the age and evolutionary history of the model species, Cyclotella nana (= Thalassiosira pseudonana). The freshwater fossil record for Thalassiosirales is extensive, well characterized, and generally supportive of a Miocene origin for the major freshwater lineages. The marine record is, by comparison, more sparse and in many cases, unverified. The discovery of freshwater thalassiosiroids in Eocene sediments pushed the freshwater fossil record considerably further back in time, highlighting an apparent gap of some 30 million years. An alternative interpretation is that the Miocene and Eocene reports represent competing hypotheses. In the absence of additional independent and decisive fossil data, I explored the relative plausibility of these two scenarios with Bayesian relaxed molecular clock methods under a range of fossil calibration schemes. Although I found no support for the Eocene fossil dates, the two major freshwater colonization events probably occurred much earlier than previously thought—as early as the Paleocene for Cyclotella, followed by an Eocene origin for the cyclostephanoid lineage. Much of the extant freshwater diversity in both lineages traces back to the Miocene, however, giving the impression of a single Miocene origin. Efforts to infer the timescale of Thalassiosirales evolution more accurately would benefit from a systematic reevaluation of the marine fossil record and formal integration of fossil species into existing phylogenetic hypotheses.

Euproops danae, a xiphosuran from the Mazon Creek (Braidwood) Fauna, had two sets of spines—genal and ophthalmic—extending from the posterior margin of its prosoma. In young individuals, the genal spines were longer than the ophthalmic spines. During growth, the relative length of each type of spine increased, but the rate of increase was much greater for ophthalmic spines. In order to explain these morphological and ontogenetic features, I have studied the hydrodynamic behavior of whole-animal models representing the actual morphology of E. danae, and various modifications thereof, at different body sizes. It can be argued that passive settling, while in an enrolled posture, was probably the primary defensive reaction that a swimming individual would have exhibited in response to an encounter with a potential predator. Experiments show that the array of spines on E. danae was an important control on the style of settling experienced by an enrolled individual. Models of the actual morphology settle steadily, while models with either longer or shorter prosomal spines tend to undergo lateral oscillations induced by turbulence in the wake. Steady fall would have rendered the horseshoe crab less perceptible to either the visual or lateral line systems of contemporary aquatic vertebrates, and thus a morphology capable of producing it would have been an important adaptation for reducing the risk of predation. This minimization of oscillatory movement would actually obtain for a variety of conceivable spine morphologies, but actual morphologies represent those ‘solutions’ which simultaneously optimize other aspects of spine function, such as mechanical protection. This optimum design changes during ontogeny because settling dynamics scale non-linearly with size. This work has both specific applications to the interpretation of similar structures in other arthropods and more general implications for the study of evolutionary functional morphology.

We have recently shown how capture-recapture models can be used in conjunction with stratigraphic range data to estimate taxonomic extinction rates and taxonomic diversity. Here we present a new method that can be used to estimate taxonomic turnover (defined here as the proportion of taxa extant at time i, that originated in the interval i – 1 to i). We used these methods in conjunction with stratigraphic range data for families in five phyla of Paleozoic marine invertebrates. We estimated fossil encounter probabilities, extinction rates, diversity, and turnover and used these estimates to test hypotheses about variation among phyla and geologic series. Encounter probabilities varied among taxa and showed evidence of a decrease over time for the geologic series examined. The number of families varied substantially among the five phyla and showed some evidence of an increase over the series examined. There was no evidence of variation in extinction probabilities among the phyla. Although there was evidence of temporal variation in extinction probabilities within phyla, there was no evidence of a linear decrease in extinction probabilities over time, as has been reported by others. We did find evidence of high extinction probabilities for the two intervals that had been identified by others as periods of mass extinction. We found no evidence of variation in turnover among the five phyla. There was evidence of temporal variation in turnover, with greater turnover occurring in the older series.

Evolutionary patterns in the articulate brachiopod hinge are investigated and this empirical example is used as a model to examine evolutionary trends and evolutionary constraints. Several features of the hinge-system geometry exhibit persistent directional change from the Cambrian to the Recent and result in increased mechanical advantage in opening the valves. These geometric changes are reflected also in general features of shell morphology and growth and in patterns of familial diversity through time. Specific, identifiable constraints on brachiopod morphology and function related to the position of the pedicle and muscles and nature of the hinge line and hinge structures may be said to direct the observed trends. The pattern of evolutionary change among all articulate brachiopods is most satisfactorily accommodated by a diffusion model of morphological change. Examined independently, the deltidiodonts (with noninterlocking hinge structures) appear to reflect a process of directional selection through time, whereas change in the cyrtomatodont hinge mechanism (with interlocking teeth and sockets) may reflect the incidental effects of selection for increasing mantle-cavity volume rather than increasing diductor (opening) muscle moment. The “transition” from deltidiodont to cyrtomatodont hinge structures over the course of brachiopod evolutionary history may be interpreted from at least two distinctly different perspectives. Paleoecologically, this transition appears to reflect a habitat shift from soft sediment to hard, rocky substrates. Phylogenetically, the transition reflects the process of evolution: deltidiodonts persist as their modified descendants, the cyrtomatodonts. Investigating the acquisition of evolutionary novelties over time, or the transition from primitive to derived morphological features, may be the most informative approach to follow in studying evolutionary trends.

The species-level properties of geographic range and geologic duration are often used as variables in evolutionary studies. However, estimates of species duration are not independent of estimates of geographic range. Before these properties are used in macroevolutionary hypotheses, error associated with these estimates must be quantified. This error may lead to spurious inferences of evolutionary processes. To assess the error associated with estimates of geographic range and geologic duration, we modeled various sampling regimes and calculated the bias associated with these estimates.

We present three analyses which document the bias associated with estimates of geographic range and geologic duration. First, we find a positive correlation between local abundance and geographic range for a sample of 180 species of Recent prosobranch gastropods from the northeastern temperate Pacific Ocean. Therefore, geographically short-ranging species are less likely to be represented in the fossil record than geographically long-ranging species because of their local rarity. Second, we demonstrate that the chance of underestimating the geographic range of a species is acute for species with restricted spatial distributions, further compounding the problem of documenting their distribution in space and time. Third, we present a simulation which quantifies the degree of autocorrelation between geographic range and geologic duration for different levels of sampling resolution and spatial distributions of fossil localities.

Data that accurately capture the spatial structure of biodiversity are required for many paleobiological questions, from assessments of changing provinciality and the role of geographic ranges in extinction and originations, to estimates of global taxonomic or morphological diversity through time. Studies of temporal changes in diversity and global biogeographic patterns have attempted to overcome fossil sampling biases through sampling standardization protocols, but such approaches must ultimately be limited by available literature and museum collections. One approach to evaluating such limits is to compare results from the fossil record with models of past diversity patterns informed by modern relationships between diversity and climatic factors. Here we use present-day patterns for marine bivalves, combined with data on the geologic ages and distributions of extant taxa, to develop a model for Pliocene diversity patterns, which is then compared with diversity patterns retrieved from the literature as compiled by the Paleobiology Database (PaleoDB). The published Pliocene bivalve data (PaleoDB) lack the first-order spatial structure required to generate the modern biogeography within the time available (<3 Myr). Instead, the published data (raw and standardized) show global diversity maxima in the Tropical West Atlantic, followed closely by a peak in the cool-temperate East Atlantic. Either today's tropical West Pacific diversity peak, double that of any other tropical region, is a purely Pleistocene phenomenon—highly unlikely given the geologic ages of extant genera and the topology of molecular phylogenies—or the paleontological literature is such a distorted sample of tropical Pliocene diversity that current sampling standardization methods cannot compensate for existing biases. A rigorous understanding of large-scale spatial and temporal diversity patterns will require new approaches that can compensate for such strong bias, presumably by drawing more fully on our understanding of the factors that underlie the deployment of diversity today.

The traditional belief that vertebrae must alternate in position with the segmented body musculature (myotomes) to allow bending of the axial skeleton is evaluated in terms of the patterns of development and structure of gnathostome vertebrae. The key functional parameter allowing lateral bending of the axial skeleton is the intersegmental position of both the neural and haemal arches, not the centrum. The intersegmental position of both the centrum and arches in tetrapods is the result of a secondary association of the centrum with the primary intersegmental position of the neural and haemal arches. The pattern of vertebral ontogeny and structure in primitive gnathostomes suggests that a causal link between sclerotomic resegmentation during amniote development and the presence of intersegmental vertebrae in the adult is spurious and corroborates the hypothesis that the process of resegmentation evolved as a method of redistributing large volumes of sclerotome cells during development. Patterns of vertebral construction in lower vertebrates are related to fast-start performance and the use of the body as a hybrid oscillator during locomotion.

Fundamental to the interpretation of bone-bearing faunal deposits is an understanding of the taphonomic processes that have modified the once living fossil community. An often neglected source of bias is that of climate-averaging, which occurs when the duration of bone accumulation exceeds the duration of an individual climatic episode. Tropical and subtropical climate change is dominated by precessional cyclicity (~21,000 year cycle), which controls monsoon rainfall intensity and thus plant communities over time. Under a climate-averaging scenario, the paleoecological characteristics of a faunal deposit represent an amalgamation of more than one phase of the precessional cycle. We investigate the degree of climate-averaging in Plio-Pleistocene bone breccias from South Africa by comparing stable isotope measurements of fossil enamel with the evidence from high-resolution speleothem paleoclimate proxies. We conclude that each of the four faunal assemblages studied are climate-averaged, having formed over a time period in excess of one-third of a precessional cycle (~7000 years). This has implications for the reconstruction of hominin paleoenvironments and estimates of Plio-Pleistocene biodiversity. We hypothesize that climate-averaging may be a common feature of tropical terrestrial vertebrate assemblages throughout the Cenozoic and Mesozoic.

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

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

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

Entelodonts are medium to large (perhaps 150–750 kg) Oligocene–Miocene bunodont artiodactyls with unique crania but typical artiodactyl postcrania. Functional and ecological interpretations are difficult because there is no clear modern analog to an entelodont in size, dentition, and cranial morphology. Entelodont crania combine primitive and derived features, including laterally expanded zygomatic arch/large temporal fossa (suggesting a large temporalis muscle), unreduced dental formula, long premolar row, fused mandibular symphysis, isognathy, and subcylindrical dentary condyles. Furthermore, the cranium has unique architectural and ontogenetic aspects unparalleled in extant mammals. Canines show heavy, carnivoran-like, apical wear in old individuals, suggesting regular contact with food. Conical premolars and associated diastemata dominate the tooth row. Premolars are often apically worn, in a fashion somewhat like those of carnivorans (e.g., Crocuta, Borophagus). Molars are low-cusped crushing teeth. Wide gape, indicated by the form of the coronoid process and temporal fossa, facilitated canine and premolar use, probably both in feeding and in social behavior. Jaw mechanics, tooth morphology, and tooth wear are compatible with omnivory and probable scavenging, an intriguing proposition for the huge Dinohyus hollandi.

Morphological responses of nonmammalian herbivores to external ecological drivers have not been quantified over extended timescales. Herbivorous nonavian dinosaurs are an ideal group to test for such responses, because they dominated terrestrial ecosystems for more than 155 Myr and included the largest herbivores that ever existed. The radiation of dinosaurs was punctuated by several ecologically important events, including extinctions at the Triassic/Jurassic (Tr/J) and Jurassic/Cretaceous (J/K) boundaries, the decline of cycadophytes, and the origin of angiosperms, all of which may have had profound consequences for herbivore communities. Here we present the first analysis of morphological and biomechanical disparity for sauropodomorph and ornithischian dinosaurs in order to investigate patterns of jaw shape and function through time. We find that morphological and biomechanical mandibular disparity are decoupled: mandibular shape disparity follows taxonomic diversity, with a steady increase through the Mesozoic. By contrast, biomechanical disparity builds to a peak in the Late Jurassic that corresponds to increased functional variation among sauropods. The reduction in biomechanical disparity following this peak coincides with the J/K extinction, the associated loss of sauropod and stegosaur diversity, and the decline of cycadophytes. We find no specific correspondence between biomechanical disparity and the proliferation of angiosperms. Continual ecological and functional replacement of pre-existing taxa accounts for disparity patterns through much of the Cretaceous, with the exception of several unique groups, such as psittacosaurids that are never replaced in their biomechanical or morphological profiles.

Application of multivariate statistical techniques, especially correspondence analysis, results in the recognition of four major communities for the Creede plant assemblages: fir-spruce forest, fir-pine forest, pine-juniper forest or woodland, and mountain mahogany chaparral. Physiognomy of the Creede assemblages indicates a mean annual temperature of <2.5°C. Paleoaltitudinal estimates based on this temperature estimate are inconsistent with physiographic and structural data of post-depositional uplift, unless the climatic effects of a closed basin are factored in; these effects imply that the Creede paleotemperature (and hence paleoaltitude) is applicable to the caldera rim. A variety of data indicate that summers were dry and most precipitation occurred as snow. The stratigraphic sequence of assemblages indicates that significant precipitational change occurred during deposition of the Creede plant-bearing beds. Chamaebatiaria, a shrub that now inhabits dry, open environments, had an ancestral taxon characteristic of the Creede fir-forest communities and strongly indicates a major shift in habitat for this lineage during the Neogene. A second genus, Luetkea, which is now herbaceous, is represented by a probable woody ancestral taxon that was common in the Creede forest communities. Consideration of the adaptive and morphologic histories of the Creede lineages suggests that physiologic adaptation may precede morphologic change. The Creede forest communities have no modern homologues.

Tyrannosaurus rex and other tyrannosaurid theropods exerted high bite forces, and large muscle attachments suggest that the tyrannosaurid neck was a concomitantly powerful component of the feeding apparatus. We examine accelerative and work-generating capacity (WGC) of neck muscles in adult Tyrannosaurus rex, using a 3-D vector-based method that incorporates aspects of muscle force generation, reconstruction of muscle morphology and moment arms, and rotational inertias of the head and neck. Under conservative assumptions, radial accelerations of the head by large superficial muscles (M. transversospinalis capitis, M. complexus, and M. longissimus capitis superficialis) enabled rapid gaze shifts and imparted high tangential velocities to food sufficient for inertial feeding. High WGC by these and deeper muscles under eccentric contraction indicate high efficacy for tearing flesh, especially with the head and neck in an extended posture. Sensitivity analyses suggest that assigned density of the antorbital region has substantial effects on calculated rotational inertia, and hence on the accuracy of results. However, even with high latitude for estimation errors, the results indicate that adult T. rex could strike rapidly at prey and engage in complexly modulated inertial feeding, as seen in extant archosaurs.

Over the past quarter-century there has been considerable innovation in methods for assessing the tempo and mode of evolution in paleobiological data sets. The current literature of these methods centers on three competing hypotheses—stasis, random walk, and directional trend—corresponding to an increasing scaling of variance with time interval (unchanging, for stasis; linear, for random walk; quadratic, for trend). For applications to a single trait there are powerful methods for discriminating among these hypotheses; but for multivariate data sets, especially the very high-dimensional multivariate data arising in image-feature-based and morphometric studies, current statistical approaches appear to be of less help. This paper proves that in the limiting case of high-dimensional morphospaces, the principal component or principal coordinate ordination of every sufficiently lengthy isotropic random walk tends to the same geometrical shape, which is not that of an ellipsoid and for which the principal components or coordinates are not independent even though they are uncorrelated. Specifically, the “scatter” of PC1 against PC2 is just a parabolic curve. The quantitative characteristics of this specific shape are not described appropriately by the corresponding “covariance structure” or Gaussian model, and the discrepancy may be pertinent to much of the existing literature of methods for differentiating among those three models of evolutionary multivariate time series. From a close examination of this common geometry of the ideal random walk model as seen in its principal components, I suggest a test for stasis, along with a mixed model illustrated by a reanalysis of some data of Gunz et al., and a related test for directional trend. These comments are intended to apply to all high-dimensional morphospaces, not just those arising in geometric morphometrics. Applications of principal components in this context distort high-dimensional data in ways that have a tendency to mislead; but these distortions can be intercepted so that studies of tempo and mode can nevertheless proceed.

Liquid removal rates in chambers of the living cephalopod Nautilus depend on two factors: the surface area of siphuncular epithelium exposed to liquid in the chambers and the buoyancy of the specimen. In Nautilus made artificially heavy, liquid removal rates are increased as much as five times above neutral buoyancy rates in surface aquaria. Although chamber volume increases faster than siphuncular surface area in successively larger chambers, the increase is small, especially in the last formed dozen chambers. Under conditions of constant liquid removal per unit area of siphuncular epithelium, this means that removal rates of liquid from any two or three successive chambers are nearly constant.

Nautilus shows ranges of chamber volume/siphuncular surface area ratios of .04 cc/mm2 (early, small chambers) to .12–.14 cc/mm2 in late, large chambers. These values are not characteristic of all chambered cephalopods. Comparison of extinct chambered cephalopod taxa with Nautilus indicate a variety of volume/area ratios for similarly sized chambers.

Plant-insect interactions are vital for structuring terrestrial ecosystems. It is still unclear how climate change in geological time might have shaped plant-insect interactions leading to modern ecosystems. We investigated the effect of Quaternary climate change on plant-insect interactions by observing insect herbivory on leaves of an evergreen sclerophyllous oak lineage (Quercus section Heterobalanus, HET) from a late Pliocene flora and eight living forests in southwestern China. Among the modern HET populations investigated, the damage diversity tends to be higher in warmer and wetter climates. Even though the climate of the fossil flora was warmer and wetter than modern sample sites, the damage diversity is lower in the fossil flora than in modern HET populations. Eleven out of 18 damage types in modern HET populations are observed in the fossil flora. All damage types in the fossil flora, except for one distinctive gall type, are found in modern HET populations. These results indicate that Quaternary climate change did not cause extensive extinction of insect herbivores in HET forests. The accumulation of a more diverse herbivore fauna over time supports the view of plant species as evolutionary “islands” for colonization and turnover of insect species.

The compound eyes of trilobites provide the best examples of fossilized sensory organs for which the function in life can be worked out today because the optical array of their corneal lenses preserves the geometry with which the eye originally sampled the visual world. An analysis of trilobite vision is strengthened by the use of new mathematical approaches to compound eye design. In particular, the product of the facet diameter (D) and the interommatidial angle (Δϕ) gives the value of the eye parameter, DΔϕ, which is a reliable indicator of the photic conditions in which the eye was used. In modern arthropods, DΔϕ values range from 0.3 for animals active in bright sunlight to 20 or more for nocturnal or deep-sea animals. Two major types of compound eyes existed in trilobites: schizochroal and holochroal. In our previous work with schizochroal eyes in the phacopids Phacops rana crassituberculata and Phacops rana milleri, we found that eye parameter values ranged from 10 to >150. These values of the eye parameter are much greater than in any living arthropod, implying that modern compound eye theory does not apply to schizochroal eyes. We suggested that each ommatidium of the schizochroal eye served as a miniature lens eye. If so, phacopid vision must have been unique, with multiply overlapping visual fields. In the new work of this paper, we examined holochroal compound eyes in Asaphus cornutus, Isotelus gigas, and Homotelus sp. Holochroal eyes contain far more ommatidia than do schizochroal types, reducing both facet diameter (D) and interommatidial angle (Δϕ). Thus, DΔϕ values in these species fall into the same range as in modern nocturnal compound eyes. This implies that function of the holochroal eye was similar to that of modern arthropods, and that they were used in moderate to dim intensities of light.

This work shows the potential for applying three-dimensional biometry to studying cell growth in larger benthic foraminifera. The volume of each test chamber was measured from the three-dimensional model obtained by means of computed tomography. Analyses of cell growth based on the sequence of chamber volumes revealed constant and significant oscillations for all investigated specimens, characterized by periods of approximately 15, 30, 90, and 360 days. Possible explanations for these periods are connected to tides, lunar cycles, and seasonality. The potential to record environmental oscillations or fluctuations during the lifetime of larger foraminifera is pivotal for reconstructing short-term paleoenvironmental variations or for gaining insight into the influence of tides or tidal current on the shallow-water benthic fauna in both recent and fossil environments.