Although generally considered rare in gastropods, septation has long been noted in turritellids, but functional hypotheses do not survive strong scrutiny. Here we outline a methodology for testing spandrel hypotheses and apply it to the problem of turritellid septa. We follow Gould in using “spandrel” as a term for all features that are nonadaptive sequelae of adaptive features of organisms, including those that are structurally necessary, those that are developmentally correlated, and nondeterministic by-products that are correlated to features under selection.

In turritellids, septa are constructed in microstructural continuity with secondary internal thickening of the shell, are highly variable features infraspecifically, and are strongly associated with degree of shell thickening. We therefore conclude that rather than being themselves adaptive, turritellid septa are spandrels of shell thickening. Turritellid septa are composed of crossed lamellar aragonite, which appears to be constructed by mantle epithelium over the visceral mass. Septation was also found in 22 of 24 gastropod families examined from a broad phylogenetic distribution. Septa thus appear to be a widespread feature of caenogastropods, in strong contrast to previous assertions that septa are less common in modern or high-spired shells.

High levels of biodiversity and endemism in ancient lakes have motivated research on evolutionary processes in these systems. Drill-core records from Lake Titicaca (Bolivia, Peru), an ancient lake in the high-elevation Altiplano, record the history of climate, landscape dynamics, and diatom evolution. That record was used to examine the patterns and drivers of morphological evolution of an endemic species complex of diatoms in the lake, the Cyclostephanos andinus complex. In an attempt to delineate species within the complex based on morphology, no discernible evidence was found for species separation based on an ordination analysis of multiple characters, but multiple populations were detected based on the distribution of valve size in individual samples. Likelihood modeling of phyletic evolution showed that size evolved through punctuated change. Correlation of size trends with environmental variables indicates that C. andinus size responded to regional environmental change driven by global processes that influenced Lake Titicaca by affecting lake level and thermal stratification.

Conodont fossils are highly valuable for Paleozoic biostratigraphy and for interpreting evolutionary change, but identifying and describing conodont morphologies, and characterizing gradual shape variation remain challenging. We used geometric morphometric (GM) analysis to conduct the first landmark-based morphometric analysis of the biostratigraphically useful conodont genus Neognathodus. Our objective is to assess whether previously defined morphotype groups are reliably distinct from one another. As such, we reevaluate patterns of morphologic change in Neognathodus P1elements, perform maximum-likelihood tests of evolutionary modes, and construct novel, GM-based biozonations through a Desmoinesian (Middle Pennsylvanian) section in the Illinois Basin. Our GM results record the entire spectrum of shape variability among Neognathodus morphotypes, thus alleviating the problem of documenting and classifying gradual morphologic transitions between morphotypes. Statistically distinct GM groups support previously established classifications of N. bassleri, N. bothrops, and N. roundyi. Statistically indistinct pairs of GM groups do not support literature designations of N. medadultimus and N. medexultimus, and N. dilatus and N. metanodosus, and we synonymize each pair. Maximum-likelihood tests of evolutionary modes provide the first statistical assessment of Neognathodus evolutionary models in the Desmoinesian. The most likely evolutionary models are an unbiased random walk or a general random walk. We name four distinct biozones through the Desmoinesian using GM results, and these align with previous biozonation structure based on the Neognathodus Index (NI), illustrating that Neognathodus-based biostratigraphic correlations would not change between GM or NI methods. The structural similarity between both biozonations showcases that determining GM-based biozones is not redundant, as this comparison validates using landmark-based GM work to construct viable biozonations for subsequent stratigraphic correlations. Although this study is limited to the Illinois Basin, our quantitative methodology can be applied broadly to test taxonomic designations of additional genera, interpret statistically robust evolutionary patterns, and construct valid biozones for this significant chordate group.

Recent studies have shown that modes of evolution, namely directional trend, random walk, and stasis, vary across morphologic traits and over the geographic range of a taxon. If so, is it possible that our interpretation of evolutionary modes is actually driven by our selection of traits in a study? In an attempt to answer this question, we have restudied the middle Miocene planktonic foraminifera Fohsella lineage, an iconic example of gradual morphologic evolution. In contrast to previous studies that have focused on the gross morphology as embodied by the edge view of tests, we analyze here multiple phenotypic traits chosen because their biologic and ecologic significance is well understood in living populations. We find that traits in the lineage did not evolve in concert. The timing and geographic pattern of changes in shape, coiling direction, size, and ecology were different. The evolution of this lineage is a mosaic combination of different evolutionary modes for different traits. We suggest that overemphasis on the evolution of some single trait, such as the edge-view outline, from narrow geographic ranges has significantly underestimated the dynamic evolutionary history of this group.

We report a detailed study of the different categories and types of abnormal morphologies in planktic foraminifera recognizable in the lowermost Danian, mainly from the El Kef and Aïn Settara sections, Tunisia. Various types of abnormalities in the test morphology were identified, including protuberances near the proloculus, abnormal chambers, double or twinned ultimate chambers, multiple ultimate chambers, abnormal apertures, distortion in test coiling, morphologically abnormal tests, attached twins or double tests, and general monstrosities. Detailed biostratigraphic and quantitative studies of the Tunisian sections documented a major proliferation of aberrant planktic foraminifera (between approximately 5% and 18% in relative abundance) during the first 200 Kyr of the Danian, starting immediately after the Cretaceous/Paleogene (K/Pg) boundary mass extinction (spanning from the Guembelitria cretacea Zone to the lower part of the P. pseudobulloides Zone). This contrasts with the proportionately low frequency of aberrant tests (generally <2%) identified within the uppermost Maastrichtian, suggesting more stable environmental conditions during the last ~50–100 Kyr of the Cretaceous. Two main pulses with abundant aberrant tests were recognized in the earliest Danian, the one recorded in the well-known K/Pg boundary clay being the more intense of those (maxima of >18%). These main pulses of aberrants coincide approximately with relevant quantitative and evolutionary turnovers in the planktic foraminiferal assemblages. In this paper, we explore the relation of these high values of the foraminiferal abnormality index with the environmental changes induced by the meteorite impact of Chicxulub in Yucatan, Mexico, and the massive eruptions of the Deccan Traps, India.

The fossil conchs of ammonoids provide valuable information about the life habits of this extinct group. A new conch measurement, the apertural surface area (ASarea), is introduced here along with modeled sizes of the buccal mass and the hyponome, based on ratios of these organs in comparison with the aperture height from the Recent Nautilus pompilius. A principal components analysis was performed using the three main characters: (1) apertural surface area index (i.e., the ratio of the apertural surface and the conch diameter), (2) buccal mass area index (i.e., the ratio between the buccal mass area and the ASarea), and (3) coiling rate of the conch. It revealed an ecomorphospace where life history traits can be tentatively assigned to species of the Ammonoidea. In this morphospace, Recent Nautilus has a marginal position, being one of the ectocochleate cephalopods with best properties for active life (capacity for handling large food items, rather good mobility). In contrast, most ammonoids possessed, at comparable conch sizes, much smaller buccal apparatuses and hyponomes, suggesting a more passive life history with reduced mobility potential and reduced capacities for larger prey items.

Ammonites have disparate adult morphologies indicative of diverse ecological niches, but ammonite hatchlings are small (~1 mm diameter), which raises questions about the similarity of egg incubation and hatchling life mode in ammonites. Modern Nautilus is sometimes used as a model organism for understanding ammonites, but despite their outward similarities, the groups are only distantly related. Trends in ammonite diversity and extinction vulnerability in the fossil record contrast starkly with those of nautilids, and embryonic shells from Late Cretaceous ammonites are two orders of magnitude smaller than nautilid embryonic shells. To investigate possible environmental changes experienced by ammonite hatchlings, we used secondary ion mass spectrometry to analyze the oxygen and carbon isotope composition of the embryonic shells and early postembryonic whorls of five juveniles of Hoploscaphites comprimus obtained from a single concretion in the Fox Hills Formation of South Dakota. Co-occurring bivalves and diagenetic calcite were also analyzed to provide a benthic baseline for comparison. The oxygen isotope ratios of embryonic shells are more like those of benthic bivalves, suggesting that ammonite eggs were laid on the bottom. Ammonite shell immediately after hatching has more negative δ18O, suggesting movement to more shallow water that is potentially warmer and/or fresher. After approximately one whorl of postembryonic growth, the values of δ18O become more positive in three of the five individuals, suggesting that these animals transitioned to a more demersal mode of life. Two other individuals transition to even lower δ18O values that could suggest movement to nearshore brackish water. These data suggest that ammonites, like many modern coleoids, may have spawned at different times of the year. Because scaphites were one of the short-term Cretaceous–Paleogene extinction survivors, it is possible that this characteristic allowed them to develop a broader geographic range and, consequently, a greater resistance to extinction.

The complex morphological evolution of the bivalve Ptychomya throughout the well-studied Agrio Formation in the Neuquén Basin (west-central Argentina, lower/upper Valanginian–lowest Barremian) constitutes an ideal opportunity to study evolutionary patterns and processes occurring at geological timescales. Ptychomya is represented in this unit by four species, the morphological variation of which needs to be temporally assessed to obtain a thorough picture of the evolution of the group. Here we use geometric morphometrics to measure variation in shell outline, ribbing pattern, and shell size in these species. We bracket the ages of our samples using a combination of ammonoid biostratigraphy and absolute ages and study the anagenetic pattern of evolution of each trait by means of paleontological time-series analysis and change tracking. We find that evolution in Ptychomya is mostly speciational, as the majority of traits show stasis, with the exceptions of shell size in P. coihuicoensis and shell outline in P. windhauseni, which seem to evolve directionally toward larger and higher shells, respectively. Ptychomya displays changes in its average morphology and disparity, which are the result of a mixture of taxonomic turnover and mosaic evolution of traits. Pulses of speciation would have been triggered by ecological opportunity, as they occur during the recovery of shallow-burrowing bivalve faunas after dysoxic events affecting the basin. On the other hand, the presence of directional patterns of evolution in P. coihuicoensis and P. windhauseni seems to be the result of a general shallowing-upward trend observed in the basin during the upper Hauterivian–lowest Barremian, as opposed to the cyclical paleoenvironmental stability inferred for the early/late Valanginian–early Hauterivian, which would have prompted stasis in P. koeneni and P. esbelta.

Intraspecific variation of organisms is of great importance to correctly carry out taxonomic work, which is a prerequisite for key disciplines in paleontology such as community paleoecology, biostratigraphy, and biogeography. However, intraspecific variation is rarely studied in ectocochleate cephalopods (ammonoids and nautiloids), for which an excessive number of taxa was established during the past centuries. Because intraspecific variation of fossilized organisms suffers from various biases (time averaging and taphonomy), an extant example is needed for actualistic comparison. We applied 3D morphometry to 93 specimens of Nautilus pompilius from three different geographic populations. This data set was used to examine the intraspecific variation throughout ontogeny in detail. Although there are slight differences between the populations as well as some measurement biases, a common pattern of intraspecific variation appears to be present. High variation in morphometric variables appears early in ontogeny and then decreases gradually in the following ontogenetic stages. Subsequently, the variation shows an increase again before maturity until a sharp increase or decrease occurs toward the end of ontogeny. Comparison with intraspecific variation of ammonoids and belemnites illustrated that some groups have ontogenetic patterns of intraspecific variation that are similar to that of N. pompilius. This implies that the abovementioned ontogenetic pattern of intraspecific variation might be common in some major cephalopod clades.

Evolving interactions between predators and prey constitute one of the major adaptive influences on marine animals during the Paleozoic. Crinoids and fish constitute a predator–prey system that may date back to at least the Silurian, as suggested by patterns of crinoid regeneration and spinosity in concert with changes in the predatory fauna. Here we present data on the frequency of breakage and regeneration in the spines of the Middle Devonian camerate Gennaeocrinus and late Paleozoic cladids, as well as an expanded survey of the prevalence of spinosity and infestation by platyceratid gastropods on crinoid genera during the Paleozoic. Spine regeneration frequency in the measured populations is comparable to arm regeneration frequencies from Mississippian Rhodocrinites and from modern deep-water crinoid populations. The prevalence of spinosity varies by taxon, time, and anatomy among Paleozoic crinoids; notably, spinosity in camerates increased from the Silurian through the Mississippian and decreased sharply during the Pennsylvanian, whereas spines were uncommon in cladids until their Late Mississippian diversification. Among camerates, tegmen spinosity is positively correlated with the presence of infesting platyceratid gastropods. These results allow us to evaluate several hypotheses for the effects of predation on morphological differences between early, middle, and late Paleozoic crinoid faunas. Our data corroborate the hypothesis that predators targeted epibionts on camerate crinoids and anal sacs on advanced cladids and suggest that the replacement of shearing predators by crushing predators after the Hangenberg extinction affected the locations of spines in Mississippian camerates.

Based on high-resolution palynological analysis of 680 samples from a core, short-term changes in plant diversity and floristic composition within the Paleogene greenhouse were detected in the lacustrine succession of a lower to middle Eocene maar lake at Messel (Federal State of Hesse, Germany). The microfloristic data show that taxonomic diversity increased rapidly within some decades during recolonization of a volcanically devastated area around the lake. With the establishment of a climax vegetation at the end of recolonization, the maximum in palynological diversity was reached within the crater area. During the following 640 Kyr the composition of the palynospectrum changed only gradually. However, different richness and evenness estimations show that alpha and gamma diversity decreased up to 35%, which can be related to the establishment of an equilibrium stage within the climax vegetation that led to the dominance of an assemblage of self-replacing species. Nevertheless, time-series analysis of alpha-diversity changes within the climax vegetation reveals that orbitally controlled climate change of Milankovitch and sub-Milankovitch order influenced the diversity of the vegetation, resulting in a rise of beta diversity. Based on the composition of the vegetation and comparison to modern analogues, our analysis proves that Eocene paratropical plant diversity increased during periods of slightly higher temperature and precipitation. Therefore, both composition and diversity of the vegetation was highly susceptible to minor-scale, short-term changes in climate, even during equable greenhouse conditions.

The placement of angiosperms and Gnetales in seed plant phylogeny remains one of the most enigmatic problems in plant evolution, with morphological analyses (which have usually included fossils) and molecular analyses pointing to very distinct topologies. Almost all morphology-based phylogenies group angiosperms with Gnetales and certain extinct seed plant lineages, while most molecular phylogenies link Gnetales with conifers. In this study, we investigate the phylogenetic signal present in published seed plant morphological data sets. We use parsimony, Bayesian inference, and maximum-likelihood approaches, combined with a number of experiments with the data, to address the morphological–molecular conflict. First, we ask whether the lack of association of Gnetales with conifers in morphological analyses is due to an absence of signal or to the presence of competing signals, and second, we compare the performance of parsimony and model-based approaches with morphological data sets. Our results imply that the grouping of Gnetales and angiosperms is largely the result of long-branch attraction (LBA), consistent across a range of methodological approaches. Thus, there is a signal for the grouping of Gnetales with conifers in morphological matrices, but it was swamped by convergence between angiosperms and Gnetales, both situated on long branches. However, this effect becomes weaker in more recent analyses, as a result of addition and critical reassessment of characters. Even when a clade including angiosperms and Gnetales is still weakly supported by parsimony, model-based approaches favor a clade of Gnetales and conifers, presumably because they are more resistant to LBA. Inclusion of fossil taxa weakens rather than strengthens support for a relationship of angiosperms and Gnetales. Our analyses finally reconcile morphology with molecules in favoring a relationship of Gnetales to conifers, and show that morphology may therefore be useful in reconstructing other aspects of the phylogenetic history of the seed plants.

Feeding behaviors may differ between past and current predators due to differences in the environments inhabited by these species at different times. We provide an example of this behavioral variability in spotted hyena (Crocuta crocuta), for which our analysis of a late Pleistocene micromammal assemblage indicates that hyenas preyed upon small rodents, a feeding habit that is rarely observed today among hyenas.

The Bois Roche cave site is situated at the edge of a low bluff overlooking the floodplain of a small stream in Cherves-Richemont (Charente, France). The deposits are dated by electron spin resonance (ESR) to about 69.7 ± 4.1 Ka. Excavations at the site recovered fossil bones and teeth of large and small mammals, together with hyena coprolites. Water screening of the sediments produced large accumulations of rodent remains with low taxonomic diversity. Small mammal bones were recovered from hyena coprolites as well. Descriptions of small mammal bone modification, both from the sediments and coprolites, are reported here. The analysis yielded a distinct taphonomic pattern representative of large carnivores (over 30 kg), which differs from any other modern or fossil predator-accumulated microfaunal assemblage taphonomically analyzed to date. To our knowledge, previous studies of hyena diet have not recorded high concentrations of a single-rodent prey species. We conclude that the low species diversity of this small mammal assemblage most likely relates to a local abundance of the prey species due to an outbreak in the rodent population, rather than from specialist predator behavior and hunting technique.

Energy availability influences natural selection on the ontogenetic histories of organisms. However, it remains unclear whether physiological controls on size remain constant throughout ontogeny or instead shift as organisms grow larger. Benthic foraminifera provide an opportunity to quantify and interpret the physicochemical controls on both initial (proloculus) and adult volumes across broad environmental gradients using first principles of cell physiology. Here, we measured proloculus and adult test dimensions of 129 modern rotaliid species from published images of holotype specimens, using holotype size to represent the maximum size of all species’ occurrences across the North American continental margin. We merged size data with mean annual temperature, dissolved oxygen concentration, particulate organic carbon flux, and seawater calcite saturation for 718 unique localities to quantify the relationship between physicochemical variables and among-species adult/proloculus size ratios. We find that correlation of community mean adult/proloculus size ratios with environmental parameters reflects covariation of adult test volume with environmental conditions. Among-species proloculus sizes do not covary identifiably with environmental conditions, consistent with the expectation that environmental constraints on organism size impose stronger selective pressures on adult forms due to lower surface area-to-volume ratios at larger sizes. Among-species adult/proloculus size ratios of foraminifera occurring in resource-limited environments are constrained by the limiting resource in addition to temperature. Identified limiting resources are food in oligotrophic waters and oxygen in oxygen minimum zones. Because among-species variations in adult/proloculus size ratios from the North American continental margin are primarily driven by the local environment’s influence on adult sizes, the evolution of foraminiferal sizes over the Phanerozoic may have been strongly influenced by changing oceanographic conditions. Furthermore, lack of correspondence between among-species proloculus sizes and environmental conditions suggests that offspring sizes in foraminifera are rarely limited by physiological constraints and are more susceptible to selection related to other aspects of fitness.

Graphoglyptids are biogenic structures commonly found in deep-sea flysch deposits and occasionally detected on the modern deep-sea floor. They extend principally horizontally and take a variety of geometric patterns, whose functional morphology remains an enigma in ichnology and paleoceanography. Based on published materials from 1850 to 2017 (79 ichnotaxa from 28 ichnogenera of graphoglyptids) and systematic observations of one of the largest deep-sea trace fossil collections in the world, this paper proposes that topological analysis is an important ingredient in the taxonomy and functional interpretation of graphoglyptids. Accordingly, graphoglyptids are classified into line, tree, and net forms by their key topological architecture, and are further attributed to 19 topological prototypes by detailed secondary topological features. Line graphoglyptids are single-connected structures with uniform tunnel width, representing primarily the feeding patterns of solitary animals. Tree graphoglyptids, the most diverse architectural group of graphoglyptids, are ascribed to 11 topological prototypes according to the connectivity features of burrow segments and the number and distributional pattern of the branching points. Net graphoglyptids are subdivided into three topological prototypes on the basis of the connectivity features and/or the regularity of the meshes. Multiconnected net forms are considered as a continuous morphological spectrum with different levels of complexity in the net formation. The various connected components in multiconnected tree and net graphoglyptids generally exhibit small and uniform tunnel diameter in a given structure (suggesting a tiny trace maker[s]). The whole structure shows relatively extensive linear or surface coverage and overall good preservation, indicating sustained processes of burrow construction. It is highly probable that certain multiconnected tree and net graphoglyptids represent some emergent patterns from self-organized collective behaviors of conspecific animals. Graphoglyptids thus provide us with a new perspective on the study of solitary and collective behaviors of macrobenthos in the deep-sea environment.

Although we do not know the cause of death of most fossil animals, mortality is often associated with ecological stress due to seasonality and other stochastic events (droughts, storms, volcanism) that may have caused shifts in feeding ecology preceding death. In these instances, dental microwear, which reflects feeding ecology in a narrow window of time, may provide a biased view of diet. Mesowear, another dental-wear proxy based on the morphology of worn cusps, requires macroscopic amounts of dental wear and reflects diet for a longer interval and may be less prone to bias from near-death ecological stress. We compared congruence between microwear and mesowear of North American, fossil rhinocerotid mass-death assemblages and collections of hunted modern rhinocerotids to test the hypothesis that fossil assemblages yield more incongruous microwear and mesowear data as a result of near-death ecological disturbances. In extant rhinos, both mesowear and microwear are associated with diet and height of the feeding environment. Mesowear and microwear in the modern rhinocerotid collections are statistically correlated, with strong relationships between average mesowear scores and labially distributed dental microwear. In contrast, a relationship between mesowear and microwear was not observed among the fossil rhinocerotid assemblages. Mesowear suggests that the fossil rhinos had low-abrasion diets, suggesting that they fed from clean, possibly tall vegetation. Some, but not all, mass-death assemblages produce microwear data with excessive scratches and/or pits compared with expectations based on mesowear results, suggesting that dental microwear was altered shortly before death in some but not all of the fossil assemblages. The dental-wear proxies available to paleoecologists provide a mosaic of dietary evidence reflecting diet over long (mesowear) and more abbreviated (microwear) periods of time that, together, provide a richer understanding of feeding ecology and its relationship to environment, seasonal change, and other ecological disturbances.

Naturally time-averaged accumulations of skeletal remains—death assemblages—provide reliable, albeit temporally coarse, information on the species composition and structure of communities in diverse settings, and their mismatch with local living communities usually signals recent human-driven ecological change. Here, we present the first test of live–dead mismatch as an indicator of human stress using ostracodes. On three islands along a gradient of human population density in the Bahamas, we compared the similarity of living and death assemblages in 10 lakes with relatively low levels of human stress to live–dead similarity in 11 physically comparable lakes subject to industrial, agricultural, or other human activities currently or in the past. We find that live–dead agreement in pristine lakes is consistently excellent, boding well for using death assemblages in modern-day and paleolimnological biodiversity assessments. In most comparison of physically similar paired lakes, sample-level live–dead mismatch in both taxonomic composition and species’ rank abundance is on average significantly greater in the stressed lakes; live–dead agreement is not lower in all samples from stressed lakes, but is more variable. When samples are pooled for lake-level and island-level comparisons, stressed lakes still yield lower live–dead agreement, but the significance of the difference with pristine lakes decreases—species that occur dead-only (or alive-only) in one sample are likely to occur alive (or dead) in other samples. Interisland differences in live–dead agreement are congruent with, but not significantly correlated with, differences in human population density. This situation arises from heterogeneity in the timing and magnitudes of stresses and in the extent of poststress recovery. Live–dead mismatch in ostracode assemblages thus may be a reliable indicator of human impact at the sample level with the potential to be a widely applicable tool for identifying impacted habitats and, perhaps, monitoring the progress of their recovery.

Lithification, the transition of unconsolidated sediments to fully indurated rocks, can potentially bias estimates of species richness, evenness, and body size distribution derived from fossil assemblages. Fossil collections made from well-indurated rocks consistently exhibit lower species richness, lower evenness, and larger average specimen size relative to collections made from unconsolidated sediments, even when collections are drawn from the same assemblage. This phenomenon is known as “lithification bias.” While the bias itself has been demonstrated empirically, much less attention has been paid to its causes. Proposed causes include taphonomic processes (e.g., destruction of small specimens during early diagenesis) and methodological differences (e.g., sieving vs. counting specimens on outcrops, bedding surfaces, or mechanically split surfaces). Here we investigate the potential effects of preferential intersection that could also result in a methodologically related bias: the preferential sampling of larger specimens relative to smaller ones when fossils are counted on rock surfaces. We used an analogue model to simulate preferential intersection (fossil collection via splitting fossiliferous rocks) and compare the results with a random-draw model that approximates the effects of sieving. The model was parameterized using nine different combinations of species abundance and species size distributions. The results show that, with rare exceptions, species richness is 5–23% lower, evenness 5–25% lower, and average specimen size 24–150% larger in preferential-intersection than in random-draw simulations. We conclude that preferential intersection can impose a significant bias independent of other mechanisms (e.g., preferential destruction of smaller specimens during diagenetic or sampling processes), that the magnitude of this bias is partially dependent on the species abundance and size distributions, and that this bias alone does not fully account for empirically observed lithification bias on species richness (i.e., other sources of bias are also at work).

Death assemblages that occupy the upper tens of centimeters of sediment in shallow-marine settings are often subject to extensive mixing, thereby limiting their usefulness in assessing environmentally mediated compositional changes through time in the local biota. Here, we provide evidence that dense, Thalassia-rich seagrass beds preserve a stratigraphic record of biotic variation because their dense root–rhizome mats inhibit mixing. We sampled benthic mollusk assemblages at seven localities in Thalassia-rich beds around St. Croix, USVI, collecting three separate sediment intervals of ~13 cm each to a total depth of ~40 cm below the sediment–water interface, and found evidence that sedimentary intervals preserved compositional stratigraphy. Further, some localities displayed systematic, directional changes down-core. An examination of interval-to-interval changes in composition revealed that compositional variation was unique from locality to locality rather than reflecting coordinated, island-wide transitions. In general, however, relative abundances of epifaunal gastropods and small lucinid bivalves tended to decrease with depth below the sediment–water interface. Quantitative comparisons of life-to-death assemblages from each successive sedimentary interval demonstrated that the shallowest death assemblages were typically more similar to the life assemblages than were deeper assemblages, suggesting that deeper intervals provide records of earlier community states.

Ecophenotypic variation in populations is driven by differences in environmental variables. In marine environments, ecophenotypic variation may be caused by differences in hydrodynamic conditions, substrate type, water depth, temperature, salinity, oxygen concentration, and habitat heterogeneity, among others. Instances of ecophenotypic variation in modern and fossil settings are common, but little is known about the influences of time averaging and spatial averaging on their preservation. Here we examine the shell morphology of two adjacent populations, both live collected and death assemblages, of the infaunal, suspension-feeding, intertidal bivalve Leukoma staminea from the well-studied Argyle Creek and Argyle Lagoon locations on San Juan Island, Washington. Individuals in the low-energy lagoon are free to burrow in the fine-grained substrate, while clams in the high-energy creek are precluded from burrowing in the rocky channel. Our results demonstrate variation in size and shape between the adjacent habitats. Lagoon clams are larger, more disk-shaped, and have relatively larger siphons than their creek counterparts, which are smaller, more spherical in shape, and have a relatively shallower pallial sinus. This ecophenotypy is preserved among death assemblages, although with generally greater variation due to time averaging and shell transport. Our interpretation is that ecophenotypic variation, in this case, is induced by differing hydrodynamic regimes and substrate types, cumulatively resulting in physiological trade-offs diverting resources from feeding and respiration to stability and shell strength, all of which have the potential to be preserved in the fossil record.