ALBERT

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

Description: Interpreting changes in ecosystem structure from the fossil record can be challenging. In a prominent example, the traditional view that brachiopods were ecologically dominant over bivalves in the Paleozoic has been disputed on both taphonomic and metabolic grounds. Aragonitic bivalves may be underrepresented in many fossil assemblages due to preferential dissolution. Abundance counts may further understate the ecological importance of bivalves, which tend to have more biomass and higher metabolic rates than brachiopods. We evaluate the relative importance of the two clades in exceptionally preserved, bulk-sampled fossil assemblages from the Pennsylvanian Breathitt Formation of Kentucky, where aragonitic bivalves are preserved as shells, not molds. At the regional scale, brachiopods were twice as abundant as bivalves and were collectively equivalent in biomass and energy use. Analyses of samples from the Paleobiology Database that contain abundance counts are consistent with these results and show no clear trend in the relative ecological importance of bivalves during the middle and late Paleozoic. Bivalves were probably more important in Paleozoic ecosystems than is apparent in many fossil assemblages, but they were not clearly dominant over brachiopods until after the Permian–Triassic extinction, which caused the shelly benthos to shift from bivalve and brachiopod dominated to merely bivalve dominated.

Description: Studies of extinction in the fossil record commonly involve comparisons of taxonomic extinction rates, often expressed as the percentage of taxa (e.g., families or genera) going extinct in a time interval. Such extinction rates may be influenced by factors that do not reflect the intrinsic severity of an extinction trigger. Two identical triggering events (e.g., bolide impacts, sea level changes, volcanic eruptions) could lead to different taxonomic extinction rates depending on factors specific to the time interval in which they occur, such as the susceptibility of the fauna or flora to extinction, the stability of food webs, the positions of the continents, and so on. Thus, it is possible for an extinction event with a higher taxonomic extinction rate to be caused by an intrinsically less severe trigger, compared to an event with a lower taxonomic extinction rate.Here, we isolate the effects of taxonomic susceptibility on extinction rates. Specifically, we quantify the extent to which the taxonomic extinction rate in a substage is elevated or depressed by the vulnerability to extinction of classes extant in that substage. Using a logistic regression model, we estimate that the taxonomic susceptibility of marine fauna to extinction has generally declined through the Phanerozoic, and we adjust the observed extinction rate in each substage to estimate the intrinsic extinction severity more accurately. We find that mass extinctions do not generally occur during intervals of unusually high susceptibility, although susceptibility sometimes increases in post-extinction recovery intervals. Furthermore, the susceptibility of specific animal classes to extinction is generally similar in times of background and mass extinction, providing no evidence for differing regimes of extinction selectivity. Finally, we find an inverse correlation between extinction rate within substages and the evenness of diversity of major taxonomic groups, but further analyses indicate that low evenness itself does not cause high rates of extinction.

Description: Extinction in the fossil record is most often measured by the percentage of taxa (species, genera, families, etc.) that go extinct in a certain time interval. This is a measure of taxonomic loss, but previous work has indicated that taxonomic loss may be decoupled from the ecological effects of an extinction. To understand the role extinction plays in ecological change, extinction should also be measured in terms of loss of functional diversity. This study tests whether ecological changes increase correspondingly with taxonomic changes during the Late Ordovician M4/M5 extinction, the Ordovician/Silurian mass extinction, and the Late Devonian mass extinction. All three extinctions are evaluated with regional data sets from the eastern United States. Ecological effects are measured by classifying organisms into ecological lifestyles, which are groups based on ecological function rather than evolutionary history. The taxonomic and ecological effects of each extinction are evaluated with additive diversity partitioning, detrended correspondence analysis, and relative abundance distributions. Although the largest taxonomic changes occur in the Ordovician/Silurian extinction, the largest ecological changes occur in the Late Devonian extinction. These results suggest that the ecological consequences of extinction need to be considered in addition to the taxonomic effects of extinction.

Description: Ecological ordination can reveal gradients in the species composition of fossil assemblages that can be correlated with paleoenvironmental gradients. Ordinations of simulated data sets suggest that nonmetric multidimensional scaling (NMDS) generally produces less distorted results than detrended correspondence analysis (DCA). We ordinated 113 brachiopod-dominated samples from the Frasnian (Late Devonian) Brallier, Scherr, and lower Foreknobs Formations of southwest Virginia, which represent a range of siliciclastic marine paleoenvironments. A clear environmental signal in the ordination results was obscured by (apparently) opportunistic species that occurred at high abundance in multiple environments; samples dominated by these species aggregated in ordination space regardless of paleoenvironmental provenance. After the opportunist-dominated samples were removed, NMDS revealed a gradient in species composition that was highly correlated with substrate (grain size); a second, orthogonal gradient likely reflects variation in disturbance intensity or frequency within grain-size regimes. Additional environmental or ecological factors, such as oxygenation, may also be related to the gradients. These two gradients, plus the environmental factors that controlled the occurrence of opportunistic species, explain much of the variation in assemblage composition in the fauna. In general, the composition of fossil assemblages is probably influenced by multiple paleoecological and paleoenvironmental factors, but many of these can be decomposed and analyzed.

Description: Many fossil assemblages are time-averaged, with multiple generations of organisms mixed into a single stratigraphic horizon. A time-averaged sample of a taxon should be more variable than a single-generation sample if enough morphologic change occurred during the interval of time-averaging. Time-averaging may also alter correlations between morphologic variables and obscure allometric relationships in an evolving population. To investigate these issues, we estimated the variability of six modern, single-generation samples of the bivalve Mercenaria campechiensis using Procrustes analysis and compared them with several time-averaged Pleistocene samples of M. campechiensis and M. permagna. Both the modern and the fossil samples ranged in variability, but these ranges were virtually identical. Morphology was quite stable over the hundreds to many thousands of years that passed as the assemblages accumulated, and the variabilities of the fossil samples could be used to estimate single-generation variability. At one fossil locality, the environment and paleocommunity changed partway through the collection interval; the morphology of Mercenaria appears stable above and below the transition but changes across it. This change is similar in magnitude to the differences between geographically separated modern populations, whereas temporal variation within single environmental settings is distinctly less than geographic variation. Analytical time-averaging (the mixing of fossils from different horizons) between paleocommunities increased variability slightly (but not significantly) above that found in living populations. While its constituent populations appear stable on millennial timescales, M. campechiensis has been evolutionarily static since at least the early to middle Pleistocene.

Description: Continental deposits of the Early Jurassic East Berlin Formation in Holyoke, Massachusetts, have yielded an exceptional occurrence of the ichnogenusTreptichnus. Here, burrows are preserved in full relief within thin mud laminae between layers of fine-grained, cross-bedded sandstone. We studied these burrows to evaluate whether earlier explanations of burrow morphology are applicable to allTreptichnus. Our research focused on three questions. (1) Do the HolyokeTreptichnushave significant vertical relief? (2) Does the lack of projections in some of the HolyokeTreptichnusresult from stratinomic sectioning through the bottom of the burrow? (3) Do expanded, bulbous ends of burrow segments result from sediment compaction? While addressing these questions, the Holyoke fossils were compared to syntype and topotype material ofTreptichnusfrom the Carboniferous of Indiana. The HolyokeTreptichnusdid not exhibit significant vertical relief, and the presence and absence of projections is explained by the positioning of new segments at different points along older ones. The bulbous ends of burrow segments resulted from trace-maker behavior, not sediment compaction. Drawing on the analysis of the Holyoke material, a new reconstruction is proposed that presents continentalTreptichnusas a shallow mole-tunnel-like burrow produced just below the sediment surface. This reconstruction is consistent with the morphology of RecentTreptichnus-like burrows produced by fly (dipteran) larvae, which are considered the most likely makers of the HolyokeTreptichnus.

Description: The taxonomic and ecologic composition of Earth's biota has shifted dramatically through geologic time, with some clades going extinct while others diversified. Here, we derive a metric that quantifies the change in biotic composition due to extinction or origination and show that it equals the product of extinction/origination magnitude and selectivity (variation in magnitude among groups). We also define metrics that describe the extent to which a recovery (1) reinforced or reversed the effects of extinction on biotic composition and (2) changed composition in ways uncorrelated with the extinction. To demonstrate the approach, we analyzed an updated compilation of stratigraphic ranges of marine animal genera. We show that mass extinctions were not more selective than background intervals at the phylum level; rather, they tended to drive greater taxonomic change due to their higher magnitudes. Mass extinctions did not represent a separate class of events with respect to either strength of selectivity or effect. Similar observations apply to origination during recoveries from mass extinctions, and on average, extinction and origination were similarly selective and drove similar amounts of biotic change. Elevated origination during recoveries drove bursts of compositional change that varied considerably in effect. In some cases, origination partially reversed the effects of extinction, returning the biota toward the pre-extinction composition; in others, it reinforced the effects of the extinction, magnifying biotic change. Recoveries were as important as extinction events in shaping the marine biota, and their selectivity deserves systematic study alongside that of extinction.