Bryozoans are active non-phototrophic biomineralizers that precipitate their calcareous skeletons in seawater. Carbonate saturation states varied temporally and spatially in Paleozoic oceans, and we used the Bryozoan Skeletal Index (BSI) to investigate whether bryozoan calcification was controlled by seawater chemistry in Paleozoic trepostome and cryptostome bryozoans. Our results show that cryptostome bryozoan genera were influenced by ocean chemistry throughout the Paleozoic and precipitated the most calcite at lower latitudes, where carbonate saturation states are generally higher, and less in midlatitudes, where carbonate will be relatively undersaturated. Trepostome bryozoan genera show a similar but weaker trend for the Ordovician to Devonian, suggesting that, like the cryptostomes, they were unable to metabolically overcome falling saturation states and simply precipitated less robust skeletons at higher latitudes. Carboniferous to Triassic trepostomes differ, however, and show a trend toward increased calcification at higher latitudes, indicating an ability to overcome unfavorable carbonate saturation states. Analysis of Permian trepostomes at the species level indicates this is most pronounced in the Southern Hemisphere, where calcification is matched by increased feeding capacity. It is proposed that this increased feeding capacity allowed trepostomes to metabolically overcome unfavorable carbonate saturation states. The differing responses of trepostome and cryptostome bryozoans to carbonate saturation states suggest that bryozoans should not be considered as a single group in marine extinctions linked to ocean chemistry changes. Likewise, it would suggest that modern stenolaemate and gymnolaemate bryozoans should be treated separately when considering their response to modern ocean chemistry changes.

Understanding current and future biodiversity responses to changing climate is pivotal as anthropogenic climate change continues. This understanding is complicated by the multitude of available metrics to quantify dynamics and by biased sampling protocols. Here, we investigate the impact of sampling protocol strategies using a data-rich fossil record to calculate effective diversity using Hill numbers for the first time on Paleogene planktonic foraminifera. We sample 22,830 individual tests, in two different size classes, across a 7 Myr time slice of the middle Eocene featuring a major transient warming event, the middle Eocene climatic optimum (MECO; ~40 Ma), at study sites in the midlatitude North Atlantic. Using generalized additive models, we investigate community responses to climatic fluctuations. After correcting for any effects of fossil fragmentation, we show a peak in generic diversity in the early and middle stages of the MECO as well as divergent trajectories between the typical size-selected community (>180 μm) and a broader assemblage, including smaller genera (>63 μm). Assemblages featuring smaller genera are more resilient to the climatic fluctuations of the MECO than those assemblages that feature only larger genera, maintaining their community structure at the reference Hill numbers for Shannon's and Simpson's indices. These results raise fundamental questions about how communities respond to climate excursions. In addition, our results emphasize the need to design studies with the aim of collecting the most inclusive data possible to allow detection of community changes and determine which species are likely to dominate future environments.

Fossils from the deep-sea Ediacaran biotas of Newfoundland are among the oldest architecturally complex soft-bodied macroorganisms on Earth. Most organisms in the Mistaken Point–type biotas of Avalonia—particularly the fractal-branching frondose Rangeomorpha— have been traditionally interpreted as living erect within the water column during life. However, due to the scarcity of documented physical sedimentological proxies associated with fossiliferous beds, Ediacaran paleocurrents have been inferred in some instances from the preferential orientation of fronds. This calls into question the relationship between frond orientation and paleocurrents. In this study, we present an integrated approach from a newly described fossiliferous surface (the “Melrose Surface” in the Fermeuse Formation at Melrose, on the southern portion of the Catalina Dome in the Discovery UNESCO Global Geopark) combining: (1) physical sedimentological evidence for paleocurrent direction in the form of climbing ripple cross-lamination and (2) a series of statistical analyses based on modified polythetic and monothetic clustering techniques reflecting the circular nature of the recorded orientation of Fractofusus misrai specimens. This study demonstrates the reclining rheotropic mode of life of the Ediacaran rangeomorph taxon Fractofusus misrai and presents preliminary inferences suggesting a similar mode of life for Bradgatia sp. and Pectinifrons abyssalis based on qualitative evidence. These results advocate for the consideration of an alternative conceptual hypothesis for position of life of Ediacaran organisms in which they are interpreted as having lived reclined on the seafloor, in the position that they are preserved.

The hyoid apparatus is essential for underwater feeding in marine tetrapods, but it is unclear whether this complex has evolved as convergently as other traits, such as dentition or locomotion. Here we compare the ossified hyoid elements in ophthalmosaurid ichthyosaurs and odontocete cetaceans, two groups with an overall similar body shape, to understand whether the hyoid elements show any signs of convergence in the context of feeding. We examined three types of data (size, morphology, and internal bone microstructure) in ophthalmosaurid and odontocete taxa in which these elements are preserved. Our data show that ichthyosaurs never experienced a shift in feeding mode, which might indicate that their hyoid apparatus never adapted to suction feeding. Also, the internal microstructure of the two animal groups differs; where the odontocetes have an overall less compact structure, ophthalmosaurid ichthyosaurs have cancellous inner cones in an outer, more compact sheath. These differences are likely explained as biomechanical adaptations to different feeding modes. Thus, the hyoid changed less and acted more as a constraint for feeding innovation in ichthyosaurs compared with cetaceans, and through a much longer time span (more than 150 Myr).

A dramatic shift from carbonate-rich to chert-rich marine strata occurred during the Permian and is frequently attributed to the increased activity of siliceous sponges and their biosiliceous sedimentation. The first-order ecologic consequences of this transition, if any, remain opaque. We analyze fossil occurrence data from the Phosphoria Basin (western North America) to test whether the presence of siliceous sponges, which are correlated with basin-wide chert strata, influenced the recruitment of benthic fauna. Using published lithologic descriptions, we categorized fossil collections by formation, facies, and lithology and used these data to code detrended correspondence analysis and nonmetric multidimensional scaling ordinations. We also analyzed the clustering of taxa into faunal units termed biofacies.

Results from these analyses indicate that fossil collections occurring in chert and carbonate are closely associated in faunal composition and community structure. These collections preferentially occur in the inner- to mid-ramp facies, in agreement with previous studies. Although largely similar in composition, collections of chert and carbonate lithology exhibit differences in the frequency and abundance of accessory brachiopod taxa (e.g., Composita and Hustedia), possibly a result of greater biosiliceous sedimentary input.

The Cambrian saw a dramatic increase in metazoan diversity and abundance. Between-assemblage diversity (beta diversity) soared in the first three Cambrian stages, suggesting a rapid increase in the geodisparity of marine animals during the Cambrian radiation. However, it remains unclear how these changes scale up to first-order biogeographic patterns. Here we outline time-traceable provinces for marine invertebrates across the Cambrian period using a compositional network based on species-level fossil occurrence data. Results confirm an increase in regional differences of faunal composition and a decrease in by-species geographic distribution during the first three stages. We also show that general biogeography tends to be reshaped after global extinction pulses. We suggest that the abrupt biogeographic differentiation during the Cambrian radiation was controlled by a combination of tectonics, paleoclimate, and dispersal capacity changes.

The frequency of biotic invasions in modern ecosystems is increasing due to global trade moving taxa outside their native ranges and climate change facilitating establishment of taxa in previously inhospitable regions. Thus, developing a holistic understanding of biotic invasions and how they impact ecosystems over different timescales—from annual to geologic timescales—is vital. Herein we examine a geologically brief invasion event, the Clarksville Phase of the Richmondian Invasion. Prior analyses have established general ecological and evolutionary patterns across the entire Richmondian Invasion, but recent sequence stratigraphic refinement makes analysis of individual invasion pulses possible for the first time. We examine biotic change across the Clarksville Phase and identify invasion impacts on diversity, paleocommunity composition, and niche stability. Invader arrival and success was strongly linked to increased propagule pressure facilitated by sea-level changes. Invaders initially colonized deep subtidal environments and then moved offshore facilitated by rapid niche evolution during the invasion interval. Invasive taxa that attained the largest population sizes belonged to previously underutilized ecological guilds. Overall, the introduction of the invasive taxa resulted in increased diversity that was maintained into the postinvasion interval accompanied by a change in community composition in which the invaders became dominant paleocommunity members. Combined, these analyses document a biotic invasion facilitated by climate change that increased local diversity through invaders occupying underutilized ecospace and competition-related niche contraction on millennial timescales. Developing a long-term perspective to accompany shorter-term studies facilitates predicting the long-term impacts of modern invasions and creating better-informed policies and practices.

Foramina of bones are beginning to yield more information about metabolic rates and activity levels of living and extinct species. This study investigates the relationship between estimated blood flow rate to the femur and body mass among cursorial birds extending back to the Late Cretaceous. Data from fossil foramina are compared with those of extant species, revealing similar scaling relationships for all cursorial birds and supporting crown bird–like terrestrial locomotor activity. Because the perfusion rate in long bones of birds is related to the metabolic cost of microfracture repair due to stresses applied during locomotion, as it is in mammals, this study estimates absolute blood flow rates from sizes of nutrient foramina located on the femur shafts. After differences in body mass and locomotor behaviors are accounted for, femoral bone blood flow rates in extinct species are similar to those of extant cursorial birds. Femoral robustness is generally greater in aquatic flightless birds than in terrestrial flightless and ground-dwelling flighted birds, suggesting that the morphology is shaped by life-history demands. Femoral robustness also increases in larger cursorial bird taxa, probably associated with their weight redistribution following evolutionary loss of the tail, which purportedly constrains femur length, aligns it more horizontally, and necessitates increased robustness in larger species.

The Asteropyginae Delo, 1935 is a group of phacopid trilobites in the family Acastidae Delo, 1935 that has served as the focus for several studies due to their distinctive morphologies and diversity. However, despite an interest in these characteristic morphologies, there have been no studies that have examined this group using morphometric techniques. Our investigation utilized both geometric morphometric and elliptical Fourier methods to quantify the morphology of cephalic sclerites of asteropyginid specimens representing wide taxonomic sampling of the clade. We constructed a phylomorphospace that shows temporal and spatial patterns of phenotypic evolution within the framework of a novel tip-dated phylogenetic tree generated using Bayesian inference. We recovered similar patterns in disparity regardless of the morphometric approach. Both analyses illustrated a marked expansion into morphospace throughout the temporal range of the clade, peaking in disparity in the Emsian and with European taxa exhibiting the highest disparity in glabellar morphospace. Additionally, glabellar shape showed low phylogenetic signal and no major patterns in phylomorphospace. This study highlights the utility of employing different methodologies to quantitatively explore the disparity of fossil taxa. It also illustrates some of the patterns of morphological change occurring during one of the final and major evolutionary radiations within Phacopida.

A length–frequency sample (n = 295) from a fossil population of the Ordovician trilobite Triarthrus eatoni Hall, 1838, assembled and analyzed by J. L. Cisne in 1973 is here reexamined using methods of length–frequency analysis commonly used in fishery science and marine biology. Theoretical considerations and the empirical data at hand suggest that the growth of T. eatoni was not “linear,” but asymptotic, as is the growth of most Recent marine invertebrates. The parameters of the von Bertalanffy growth function (L∞ = 41 mm, K = 0.29 yr−1) suggest that T. eatoni, which apparently lived in a challenging environment, grew somewhat more slowly than the extant marine isopod Ceratoserolis trilobitoides (Eights, 1833), used here as Recent analogue to T. eatoni. This trilobite probably lived up to 10 years, rather than the suggested 4 years, and its mortality rate was 15%–20% per year rather than 30%–40% per year. These represent the first estimates of trilobite absolute growth characteristics using methods known to accurately model growth in extant water-breathing ectotherms. These provide a baseline for trilobite growth that can be used to make inferences about growth in other species. The approach used here may also be applied to other trilobites for which suitable length–frequency data exist.

Understanding spatial variation in origination and extinction can help to unravel the mechanisms underlying macroevolutionary patterns. Although methods have been developed for estimating global origination and extinction rates from the fossil record, no framework exists for applying these methods to restricted spatial regions. Here, we test the efficacy of three metrics for regional analysis, using simulated fossil occurrences. These metrics are then applied to the marine invertebrate record of the Permian and Triassic to examine variation in extinction and origination rates across latitudes. Extinction and origination rates were generally uniform across latitudes for these time intervals, including during the Capitanian and Permian–Triassic mass extinctions. The small magnitude of this variation, combined with the possibility of its attribution to sampling bias, cautions against linking any observed differences to contrasting evolutionary dynamics. Our results indicate that origination and extinction levels were more variable across clades than across latitudes.

Melanin pigments are central to colors and patterns in modern vertebrate integuments, which inform upon ecological and behavioral strategies like crypsis, aposematism, and sociosexual selection. Over the last decade, melanin has emerged as a valuable tool for predicting color in exceptionally preserved fossil feathers and subsequent testing of paleobiological hypotheses in long-extinct dinosaurs and birds. Yet much remains to be learned about melanin stability, diagenetic alterations to melanin chemistry, and their implications for “paleocolor reconstruction.” Pressure–temperature maturation experiments with modern feathers offer a way to examine these topics but have mostly been conducted in closed-system capsules or open-system aluminum foil. Both methods have operational limitations and do not consider the filtering effect of porous sediment matrices on thermally labile chemical groups versus stable ones during natural fossilization. We use sediment-encased maturation to resolve this issue and demonstrate replication of organic preservation of melanin highly comparable to compression fossils. Our experiments, coupled with time-of-flight secondary ion mass spectrometry, show predictable volatilization of N/S-bearing molecules and increased melanin cross-linking with elevated temperatures. We also suggest that eumelanin is more stable compared with pheomelanin at higher temperatures, explaining why eumelanic colors (black, dark brown, iridescent) are better preserved in fossils than pheomelanic ones (reddish brown). Furthermore, we propose that proteins preferentially undergo hydrolysis more so than forming N-heterocycles in selectively open systems analogous to natural matrices. Thus, we conclude that melanin pigments and not diagenetically altered protein remnants are the key players in promoting fossilization of soft tissues like feathers.

The rise of jawed vertebrates (gnathostomes) and extinction of nearly all jawless vertebrates (agnathans) is one of the most important transitions in vertebrate evolution, but the causes are poorly understood. Competition between agnathans and gnathostomes during the Devonian period is the most commonly hypothesized cause; however, no formal attempts to test this hypothesis have been made. Generally, competition between species increases as morphological similarity increases; therefore, this study uses the largest to date morphometric comparison of Silurian and Devonian agnathan and gnathostome groups to determine which groups were most and least likely to have competed. Five agnathan groups (Anaspida, Heterostraci, Osteostraci, Thelodonti, and Furcacaudiformes) were compared with five gnathostome groups (Acanthodii, Actinopterygii, Chondrichthyes, Placodermi, and Sarcopterygii) including taxa from most major orders. Morphological dissimilarity was measured by Gower's dissimilarity coefficient, and the differences between agnathan and gnathostome body forms across early vertebrate morphospace were compared using principal coordinate analysis. Our results indicate competition between some agnathans and gnathostomes is plausible, but not all agnathan groups were similar to gnathostomes. Furcacaudiformes (fork-tailed thelodonts) are distinct from other early vertebrate groups and the least likely to have competed with other groups.

Paleontologists have long stressed the need to know how sampling the fossil record might influence our knowledge of the evolution of life. Here, we combine fossil occurrences of North American marine invertebrates from the Paleobiology Database with lithologic data from Macrostrat to identify sampling patterns in carbonate and siliciclastic rocks. We aim to quantify temporal trends in sampling effort within and between lithologies, focusing on the proportion of total available volume that has been sampled (sampled fossiliferous proportion, here called κ). Results indicate that the sampled fossiliferous proportion was stable during the Paleozoic, and variable during the post-Paleozoic, but showed no systematic increase through time. Fossiliferous carbonate rocks are proportionally more sampled than siliciclastic rocks, with intervals where the carbonate κ is double the siliciclastic κ. Among possible explanations for the apparent oversampling of fossiliferous carbonate rocks, analyses suggest that barren units, taphonomic dissolution, or data entry errors cannot completely explain sampling patterns. Our results suggest that one of the important drivers might be that paleontologists publish taxonomic descriptions from carbonate rocks more frequently. The higher diversity in carbonate rocks might account for an ease in the description of unknown species and therefore a higher rate of published fossils. Finally, a strong effect in favor of carbonate rocks might distort our perception of diversity through time, even under commonly used standardization methods. Our results also confirm that previous descriptions of an increase in the proportion of sampled fossiliferous rocks over time were driven by the sampling of the nonmarine fossil record.

Ammonoid cephalopods were Earth's most abundant oceanic carnivores for hundreds of millions of years, yet their probable range of swimming capabilities is poorly constrained. We investigate potential hydrodynamic costs and advantages provided by different conch geometries using computational fluid dynamics simulations. Simulations of raw drag demonstrate expected increases with velocity and conch inflation, consistent with published experimental data. Analysis at different scales of water turbulence (via Reynolds number) reveals dynamic trade-offs between conch shape, size, and velocity. Among compressed shells, the cost of umbilical exposure makes little difference at small sizes (and/or low velocity) but is profound at large sizes (and/or high velocity). We estimate that small ammonoids could travel one to three diameters per second (i.e., a typical ammonoid with a 5-cm-diameter shell could travel 5–15 cm/s), but that large ammonoids faced greater discrepancies (a 10 cm serpenticone likely traveled <30 cm/s, while a 10 cm oxycone might achieve >40 cm/s). All of these velocities are proposed only for short bursts of jet propulsion, lasting only a few seconds, in the service of dodging a predator or conspecific rival. These analyses do not include phylogeny, taxonomy, second-order conch architecture (ribs, ornament, etc.), or hydrostatic consequences of internal anatomy (soft body, suture complexity). For specific paleoecological context, we consider how these results inform our reconstruction of Jurassic ammonite recovery from the end-Triassic mass extinction. Greater refinements will come with additional simulations that measure how added mass is influenced by individual shape-trait variations, ornament, and subtle body extensions during a single jet motion.

Understanding how time averaging changes during burial is essential for using Holocene and Anthropocene cores to analyze ecosystem change, given the many ways in which time averaging affects biodiversity measures. Here, we use transition-rate matrices to explore how the extent of time averaging changes downcore as shells transit through a taphonomically complex mixed layer into permanently buried historical layers: this is a null model, without any temporal changes in rates of sedimentation or bioturbation, to contrast with downcore patterns that might be produced by human activity. Assuming stochastic burial and exhumation movements of shells between increments within the mixed layer and stochastic disintegration within increments, we find that almost all combinations of net sedimentation, mixing, and disintegration produce a downcore increase in time averaging (interquartile range [IQR] of shell ages), this trend is typically associated with a decrease in kurtosis and skewness and by a shift from right-skewed to symmetrical age distributions. A downcore increase in time averaging is thus the null expectation wherever bioturbation generates an internally structured mixed layer (i.e., a surface, well-mixed layer is underlain by an incompletely mixed layer): under these conditions, shells are mixed throughout the entire mixed layer at a slower rate than they are buried below it by sedimentation. This downcore trend created by mixing is further amplified by the downcore decline in disintegration rate. We find that transition-rate matrices accurately reproduce the downcore changes in IQR, skewness, and kurtosis observed in bivalve assemblages from the southern California shelf. The right-skewed shell age-frequency distributions typical of surface death assemblages—the focus of most actualistic research—might be fossilized under exceptional conditions of episodic anoxia or sudden burial. However, such right-skewed assemblages will typically not survive transit through the surface mixed layer into subsurface historical layers: they are geologically transient. The deep-time fossil record will be dominated instead by the more time-averaged assemblages with weakly skewed age distributions that form in the lower parts of the mixed layer.

The nature of phenotypic evolution within lineages is central to many unresolved questions in paleontology and evolutionary biology. Analyses of evolutionary time series of ancestor–descendant populations in the fossil record are likely to make important contributions to many of these debates. However, the limited number of models that have been applied to these types of data may restrict our ability to interpret phenotypic evolution in the fossil record. Using uni- and multivariate models of trait evolution that make different assumptions regarding the dynamics of the adaptive landscape, I evaluate contrasting hypotheses to explain evolution of size in the radiolarian Eucyrtidium calvertense and armor in the stickleback Gasterosteus doryssus. Body-size evolution in E. calvertense is best explained by a model in which the lineage evolves as a consequence of a shift in the adaptive landscape that coincides with the initiation of neosympatry with its sister lineage. Multivariate evolution of armor traits in a stickleback lineage (G. doryssus) shows evidence of adaptation toward independent optima on the adaptive landscape at the same time as traits change in a correlated fashion. The fitted models are available in the R package evoTS, which builds on the paleoTS framework.

Biogeochemical analyses of organisms’ tissues provide direct proxies for diets, behaviors, and environmental interactions that have proven invaluable for studies of extant and extinct species. Applying these to Cretaceous ecosystems has at times produced anomalous results, however, as dinosaurs preserve unusually positive stable carbon isotope compositions relative to extant C3-feeding vertebrates. This has been hypothesized to be a unique property of dinosaur dietary physiology, with potential significance for our interpretations of their paleobiology. We test that hypothesis through multi-taxic stable carbon isotope analyses of a spatiotemporally constrained locality in the Late Cretaceous of Canada, and compare the results to a modern near-analogue environment in Louisiana. The stable carbon isotope anomaly is present in all sampled fossil vertebrates, dinosaur or not. This suggests another more widespread factor is responsible. Examinations of diagenetic effects suggest that, where present, they are insufficient to explain the isotope anomaly. The isotope anomaly is therefore not primarily the result of a unique dietary physiology of dinosaurs, but rather a mix of factors impacting all taxa, such as environmental and/or source-diet differences. Our study underscores the importance of multi-taxic samples from spatiotemporally constrained localities in testing hypotheses of extinct organisms and ecosystems, and in the use of modern data to “ground truth” when evaluating analogue versus non-analogue conditions in greenhouse paleoecosystems.

Despite the rich fossil record of Neogene chondrichthyans (chimaeras, sharks, rays, and skates) from Europe, little is known about the macroevolutionary processes that generated their current diversity and geographical distribution. We compiled 4368 Neogene occurrences comprising 102 genera, 41 families, and 12 orders from four European regions (Atlantic, Mediterranean, North Sea, and Paratethys) and evaluated their diversification trajectories and paleobiogeographic patterns. In all regions analyzed, we found that generic richness increased during the early Miocene, then decreased sharply during the middle Miocene in the Paratethys, and moderately during the late Miocene and Pliocene in the Mediterranean and North Seas. Origination rates display the most significant pulses in the early Miocene in all regions. Extinction rate pulses varied across regions, with the Paratethys displaying the most significant pulses during the late Miocene and the Mediterranean and North Seas during the late Miocene and early Pliocene. Overall, up to 27% and 56% of the European Neogene genera are now globally and regionally extinct, respectively. The observed pulses of origination and extinction in the different regions coincide with warming and cooling events that occurred during the Neogene globally and regionally. Our study reveals complex diversity dynamics of Neogene chondrichthyans from Europe and their distinct biogeographic composition despite the multiple marine passages that connected the different marine regions during this time.

We examine temporal and spatial variation in morphology of the ammonoid cephalopod Discoscaphites iris using a large dataset from multiple localities in the Late Cretaceous (Maastrichtian) of the U.S. Gulf and Atlantic Coastal Plains, spanning a distance of 2000 km along the paleoshoreline. Our results suggest that the fossil record of D. iris is consistent with no within-species net accumulation of phyletic evolutionary change across morphological traits or the lifetime of this species. Correlations between some traits and paleoenvironmental conditions as well as changes in the coefficient of variation may support limited population-scale ecophenotypic plasticity; however, where stratigraphic data are available, no directional changes in morphology occur before the Cretaceous/Paleogene (K/Pg) boundary. This is consistent with models of “dynamic” evolutionary stasis. Combined with knowledge of life-history traits and paleoecology of scaphitid ammonoids, specifically a short planktonic phase after hatching followed by transition to a nektobenthic adult stage, these data suggest that scaphitids had significant potential for rapid morphological change in conjunction with limited dispersal capacity. It is therefore likely that evolutionary mode in the Scaphitidae (and potentially across the broader ammonoid clade) follows a model of cladogenesis wherein a dynamic morphological stasis is periodically interrupted by more substantial evolutionary change at speciation events. Finally, the lack of temporal changes in our data suggest that global environmental changes had a limited effect on the morphology of ammonoid faunas during the latest Cretaceous.