The Strophomenata, which includes two large orders, the Strophomenida and Productida, is the largest group of Paleozoic brachiopods. Nearly all uncemented strophomenatans possessed an unusual concave brachial valve. Most of these have been considered to have lived epifaunally, but had they rested on the seafloor, not only would they have faced intense predation, but their physical instability would have been fatal. I conclude that nearly all strophomenatans, like similar concavo-convex pectinid bivalves, lived infaunally by ejecting water to create a pit into which they descended, to be protected by sediment covering the concave valve. Strophomenatans have been discovered with this sediment preserved in place. If exhumed and turned upside down, a strophomenatan could have righted itself by squirting water. Many productides had anchoring spines, and some had hinge areas with stabilizing flanges. Small spines on the brachial valves of some productides served to trap disguising sediment. Evolutionary loss of hinge teeth within both the Strophomenida and Productida reduced the friction of valve clapping. Partly because of their slender shape, strophomenides were typically more vulnerable to exhumation than productides. Strophomenides also ejected water less effectively than productides and would have been less adept at righting themselves. The virtual disappearance of the strophomenides during the Devonian can be attributed to their vulnerability to intensifying benthic bulldozing and predation. The success of the productides during the late Paleozoic can be attributed to their relatively deep sequestration in the sediment and ability to right themselves and reburrow effectively when exhumed and overturned.

The study of spatial patterns in biotic compositional variability in deep time is key to understanding the macroecological response of species assemblages to global change. Globally warm climatic phases are marked by the expansion of megathermal climates into currently extra-tropical areas. However, there is currently little information on whether vegetation in these “paratropical” regions resembled spatially modern tropical or extra-tropical biomes. In this paper we explore spatial heterogeneity in extra-tropical megathermal vegetation, using sporomorph (pollen and spore) data from the late Paleocene Calvert Bluff and Tuscahoma Formations of the formerly paratropical U.S. Gulf Coast (Texas, Mississippi, and Alabama). The data set comprises 139 sporomorph taxa recorded from 56 samples. Additive diversity partitioning, nonmetric multidimensional scaling, and cluster analysis show compositional heterogeneity both spatially and lithologically within the U.S. Gulf Coastal Plain (GCP) microflora. We then use sporomorph data from Holocene lake cores to compare spatial patterns in the late Paleocene GCP with modern tropical and extra-tropical biomes. Distance decay analysis of the Holocene data reveals a higher rate of spatial turnover in tropical versus extra-tropical vegetation types, consistent with a latitudinal gradient in floral compositional heterogeneity. The specific combination of rate and scale dependency of distance decay in the Holocene assemblages prevented us from associating the late Paleocene GCP with any particular modern biome. Our results demonstrate the importance of spatial scale, taphonomy, and lithology in determining patterns of spatial heterogeneity, and show the potential of the fossil sporomorph record for studying spatial patterns and processes in deep time.

Understanding the distribution of taxa in space and time is key to understanding diversity dynamics. The fossil record provides an avenue to assess these patterns on vast timescales and through major global changes. The Eublastoidea were a conservatively plated Paleozoic echinoderm clade that range from the middle Silurian to the end-Permian. The geographic distribution of the eublastoids, as a whole, has been qualitatively assessed but has historically lacked a quantitative analysis. This is the first examination of the Eublastoidea using probabilistic methods within the R package BioGeoBEARS to assess macroevolutionary trends. Results provide an updated understanding of eublastoid diversity with new peaks and troughs in diversity through their evolutionary history. Lithology is examined in an evolutionary framework and does not have clear evolutionary trends, and there is much work to be done regarding environmental preferences. Biogeographic patterns do not recover precise group origins but do support the previous work that outlines Eublastoidea as a Laurentian clade. Sympatric speciation events dominant the clade's history but are likely exaggerated due to the highly combined areas. Vicariance events are rare and restricted to the Silurian and Devonian, and dispersal events are more common throughout the evolutionary history. Pathways allowing for lineage migrations are noted between southern Laurussia and China in the Devonian and Carboniferous and southern Laurussia and eastern Gondwana in the Carboniferous. Future work will include the addition of more non-Laurentian species into the estimated phylogeny to better estimate these global patterns.

Strata of the Ediacaran Period (635–538.8 Ma) yield the oldest known fossils of complex, macroscopic organisms in the geologic record. These “Ediacaran-type” macrofossils (known as the Ediacaran biota) first appear in mid-Ediacaran strata, experience an apparent decline through the terminal Ediacaran, and directly precede the Cambrian (538.8–485.4 Ma) radiation of animals. Existing hypotheses for the origin and demise of the Ediacaran biota include: changing oceanic redox states, biotic replacement by succeeding Cambrian-type fauna, and mass extinction driven by environmental change. Few studies frame trends in Ediacaran and Cambrian macroevolution from the perspective of the sedimentary rock record, despite well-documented Phanerozoic covariation of macroevolutionary patterns and sedimentary rock quantity. Here we present a quantitative analysis of North American Ediacaran–Cambrian rock and fossil records from Macrostrat and the Paleobiology Database. Marine sedimentary rock quantity increases nearly monotonically and by more than a factor of five from the latest Ediacaran to the late Cambrian. Ediacaran–Cambrian fossil quantities exhibit a comparable trajectory and have strong (rs > 0.8) positive correlations with marine sedimentary area and volume flux at multiple temporal resolutions. Even so, Ediacaran fossil quantities are dramatically reduced in comparison to the Cambrian when normalized by the quantity of preserved marine rock. Although aspects of these results are consistent with the expectations of a simple fossil preservation–induced sampling bias, together they suggest that transgression–regression and a large expansion of marine shelf environments coincided with the diversification of animals during a dramatic transition that is starkly evident in both the sedimentary rock and fossil records.

Identifying correlates of extinction risk is important for understanding the underlying mechanisms driving differential rates of extinction and variability in the temporal durations of taxa. Increasingly, it is recognized that the effects of multiple, potentially interacting variables and phylogenetic relationships should be incorporated when studying extinction selectivity to account for covariation of traits and shared evolutionary history. Here, I explore a variety of biological and ecological controls on genus longevity in the global fossil record of diplobathrid crinoids by analyzing the combined effects of species richness, habitat preference, body size, filtration fan density, and food size selectivity. I employ a suite of taxic and phylogenetic approaches to (1) quantitatively compare and rank the relative effects of multiple factors on taxonomic longevity and (2) determine how phylogenetic comparative approaches alter interpretations of extinction selectivity.

I find controls on diplobathrid genus duration are hierarchically structured, where species richness is the primary predictor of duration, habitat is the secondary predictor, and combinations of ecological and biological traits are tertiary controls. Ecology plays an important but complex role in the generation of crinoid macroevolutionary patterns. Notably, tolerance of environmental heterogeneity promotes increased genus duration across diplobathrid crinoids, and the effects of traits related to feeding ecology vary depending on habitat lithology. Finally, I find accounting for phylogeny does not consistently decrease the significance of correlations between traits and genus duration, as is commonly expected. Instead, the strength of relationships between traits and duration may increase, decrease, or remain statistically similar, and both the magnitude and direction of these shifts are generally unpredictable. However, traits with strong correlations and/or moderately large effect sizes (Cohen's f2 > 0.15) under taxic approaches tend to remain qualitatively unchanged under phylogenetic approaches.

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).

Preferential dissolution of the biogenic carbonate polymorph aragonite promotes preservational bias in shelly marine faunas. While field studies have documented the impact of preferential aragonite dissolution on fossil molluscan diversity, its impact on regional and global biodiversity metrics is debated. Epicontinental seas are especially prone to conditions that both promote and inhibit preferential dissolution, which may result in spatially extensive zones with variable preservation. Here we present a multifaceted evaluation of aragonite dissolution within the Late Cretaceous Western Interior Seaway of North America. Occurrence data of mollusks from two time intervals (Cenomanian/Turonian boundary, early Campanian) are plotted on new high-resolution paleogeographies to assess aragonite preservation within the seaway. Fossil occurrences, diversity estimates, and sampling probabilities for calcitic and aragonitic fauna were compared in zones defined by depth and distance from the seaway margins. Apparent range sizes, which could be influenced by differential preservation potential of aragonite between separate localities, were also compared. Our results are consistent with exacerbated aragonite dissolution within specific depth zones for both time slices, with aragonitic bivalves additionally showing a statistically significant decrease in range size compared with calcitic fauna within carbonate-dominated Cenomanian–Turonian strata. However, we are unable to conclusively show that aragonite dissolution impacted diversity estimates. Therefore, while aragonite dissolution is likely to have affected the preservation of fauna in specific localities, time averaging and instantaneous preservation events preserve regional biodiversity. Our results suggest that the spatial expression of taphonomic biases should be an important consideration for paleontologists working on paleobiogeographic problems.

Stanley's (1979) method of estimating taxonomic longevity from Lyellian curves differs from Kurtén's (1960) because of different underlying assumptions. Kurtén's model supposes that taxon richness and longevity are stochastically invariate over the period used for the Lyellian estimate. Both Kurtén and Stanley seek to calculate the average longevity of a taxon, when the temporal stratigraphic ranges are known after and before a given time datum. Under the exponential decay model used by Kurtén, mean taxonomic longevity is indeed approximately 2.89 half-lives, as he maintained. Stanley's estimate is based on the unrealistic assumptions of uniform longevity and evenly-spaced origins of fossil taxa. This leads to an estimate of longevity as exactly two half-lives.

Deeper knowledge about how species and communities respond to climate change and environmental gradients should be supported by evidence from the past, especially as modern responses are influenced by anthropogenic pressures, including human population growth, habitat destruction and fragmentation, and intensifying land use. There have been great advances in modeling species’ geographic distributions over shallow time, where consideration of evolutionary change is likely less important due to shorter time for evolution and speciation to occur. Over these shallow time periods, we have more resources for paleoclimate interpretation across large geographic landscapes. We can also gain insight into species and community changes by studying deep records of temporal changes. However, modeling species geographic distributions in deep time remains challenging, because for many species there is sparse coverage of spatial and temporal occurrences and there are fewer paleoclimate general circulation models (GCMs) to help interpret the geographic distribution of climate availability. In addition, at deeper time periods, it is essential to consider evolutionary change within lineages of species. I will discuss a framework that integrates evolutionary information in the form of phylogenetic relatedness from clades of extant closely related species, where and when there are associated fossil occurrences, and the geographic distribution of paleoclimate in deep time to infer species past geographic response to climate change and to estimate where and when there were hotspots of ancient diversification. More work is needed to better understand the evolution of physiological tolerances and how physiological tolerances relate to the climate space in which species occur.

Geographically explicit, taxonomically resolved fossil occurrences are necessary for reconstructing macroevolutionary patterns and for testing a wide range of hypotheses in the Earth and life sciences. Heterogeneity in the spatial and temporal distribution of fossil occurrences in the Paleobiology Database (PBDB) is attributable to several different factors, including turnover among biological communities, socioeconomic disparities in the intensity of paleontological research, and geological controls on the distribution and fossil yield of sedimentary deposits. Here we use the intersection of global geological map data from Macrostrat and fossil collections in the PBDB to assess the extent to which the potentially fossil-bearing, surface-expressed sedimentary record has yielded fossil occurrences. We find a significant and moderately strong positive correlation between geological map area and the number of fossil occurrences. This correlation is consistent regardless of map unit age and binning protocol, except at period level; the Neogene and Quaternary have non-marine map units covering large areas and yielding fewer occurrences than expected. The sedimentary record of North America and Europe yields significantly more fossil occurrences per sedimentary area than similarly aged deposits in most of the rest of the world. However, geographic differences in area and age of sedimentary deposits lead to regionally different expectations for fossil occurrences. Using the sampling of surface-expressed sedimentary units in North America and Europe as a predictor for what might be recoverable from the surface-expressed sedimentary deposits of other regions, we find that the rest of the globe is approximately 45% as well sampled in the PBDB. Using age and area of bedrock and sampling in North America and Europe as a basis for prediction, we estimate that more than 639,000 occurrences from outside these regions would need to be added to the PBDB to achieve global geological parity in sampling. In general, new terrestrial fossil occurrences are expected to have the greatest impact on our understanding of macroevolutionary patterns.

Arthropods are known to display a variable number of eye lenses and this number mostly increases during their development. In trilobites, most species possessing schizochroal eyes exhibit a notable intraspecific variation in the number of dorso ventral files of eye lenses that can be age related (i.e., growth) or not (i.e., living environment). Several previous studies have shown that some trilobite groups (e.g. phacopids) tend to have fewer lenses/files in representatives from the deeper habitats than those from shallower habitats. In this study, we analyzed the pattern of variation in the number of dorso ventral files of eye lenses in two Devonian phacopid trilobites from the Prague Basin of the Czech Republic. We quantified their intraspecific variability. To better understand the patterning, we compared more than 120 individuals. Data first reveal evidence of a bimodal distribution of lens/file number without intermediate forms among each of two studied populations of Prokops prokopi (Chlupáč, 1971) and throughout the ontogeny of Pedinopariops insequens (Chlupáč, 1977). Our results indicate that caution must be taken for taxonomical affiliation and biodiversity analyses of taxa in which the intraspecific variability is unclear. Additionally, we investigated possible relations of these bimodalities to the stratigraphical position of studied populations and to the paleoenvironment. In Prokops prokopi, a slightly different age of both populations, together with supposed differences in the local environments can be responsible for observed variability. In Pedinopariops insequens, stress conditions possibly related to the approaching onset of the Basal Choteč Event can be responsible for surprising intrapopulation variability. We speculate that the stress conditions could cause a bimodal selection and possibly also the change of ontogenetic trajectory within this species. Pedinopariops insequens was the only phacopid in the Prague Basin that crosses the Lower/Middle Devonian boundary and survived also the onset of Basal Choteč Event.

Colony-wide feeding currents are a common feature of many bryozoan colonies. These feeding currents are centered on excurrent macular chimneys that expel previously filtered water away from the colony surface. In some bryozoans these macular chimneys consist of a branching channel network that converges at a point in the center of the chimney. The bifurcating channels of the maculae are analogous to a stream channel network in a closed basin with centripetal drainage. The classical methods of stream channel network analysis from geomorphology are here used to quantitatively analyze the number and length of macular channels in bryozoans. This approach is applied to a giant branch of the trepostome bryozoan Tabulipora from the Early Permian Kim Fjelde Formation in North Greenland. Its large size allowed 18 serial tangential peels to be made through the 8-mm-thick exozone. The peels intersected two stellate maculae as defined by contiguous exilapores. The lengths of 1460 channels radiating from the maculae were measured and their Horton-Strahler stream order and Shreve magnitude scored.

We hypothesize that if fossil bryozoan maculae function as excurrent water chimneys, then they should conform to Horton's laws of stream networks and behave like closed basins with centripetal drainage. Results indicate that the stellate maculae in this bryozoan behaved liked stream channel networks exhibiting landscape maturation and stream capture. They conformed to the Law of Stream Number. They have a Bifurcation Ratio that falls within the range of natural stream channel networks. They showed a pattern opposite that expected by the Law of Stream Lengths in response to behavior characteristic of a centripetal drainage pattern in a closed basin. Thus, the stellate maculae in this bryozoan probably functioned as excurrent water chimneys with the radiating channels serving to efficiently collect the previously filtered water, conducting it to the central chimney for expulsion away from the colony surface.

New growth rate estimates for nine species from three genera of New Zealand Crassatellidae (Mollusca: Bivalvia), combined with existing morphometric ontogenetic descriptions, allow identification of heterochronic processes in the evolution of these genera. Both paedomorphosis (progenesis and neoteny) and peramorphosis (hypermorphosis and acceleration) have occurred within the clade. Overall, morphological variability and response to environmental pressure in this nonsiphonate group is restricted by the interplay of anatomical and life habit constraints. Stability in the substrate, predator avoidance, sluggish burrowing speed, and inability to escape by deep burial are suggested as key drivers of, or constraints on, morphological change. Two groups of shell characters are identified: heavy, armored “anchors” and elongate “snorkels,” which combine juvenile and adult traits in shells of different sizes and ages, produced by heterochronic variation in developmental timing. Anchors and snorkels both represent different “solutions” to the problems of life as a nonsiphonate, infaunal bivalve.

An understanding of the thermal strategies of extant animals is a prerequisite for any useful discussion on the possible thermal strategies of dinosaurs. Of cardinal importance is the identification of the selection pressures that have resulted in the evolution of high body temperatures. Although different selection pressures have probably been operational in different organisms, enhanced muscle power may have been a prime factor.

Many dinosaurs, including the ornithomimids, carnosaurs and hadrosaurs possess cursorial features suggesting high activity levels, which in turn suggest the possession of high body temperatures. Even if it is denied that high body temperatures are a prerequisite for speed, it cannot be denied that muscular activity liberates large quantities of heat, and most dinosaurs, by virtue of their small area to volume ratio, must have maintained constant, and sometimes high, body temperatures. This thermal strategy, described as inertial homeothermy, does not require a high (mammalian/avian) rate of metabolism, and most dinosaurs were probably not endothermic.

The smallest dinosaurs (e.g. Compsognathus) were probably too small to have been able to maintain constant body temperatures in the absence of external insulation. Perhaps they were feathered, like their close relative Archaeopteryx. Presence of feathers in Archaeopteryx is compelling evidence for endothermy.

The largest dinosaurs, the sauropods, had an exceedingly small area to volume ratio, that probably imposed limitations on their rate of heat dissipation. This in turn may have restricted their rate of muscular activity; there are also sound mechanical reasons why they should have been relatively inactive. It is therefore concluded that sauropods probably maintained relatively low body temperatures.

Biostratigraphic applications of Devonian plant macrofossils are unusual because of the rarity of these fossil plants. Moreover, the taxonomic determination of these plants is too often either arguable or impossible. Here is proposed a biostratigraphic method based on a quantification of individual plant characters and not on determination of whole fossils. The method is the following: (1) each biocharacter found at a given locality receives a score according to whether it is primitive or derived, so that the biocharacter score increases with increasing derivation; (2) the biostratigraphic coefficients of well-dated localities are calculated, the biostratigraphic coefficient of a given plant locality being the mean of all the biocharacter scores of this locality; (3) the scores of these well-dated localities are used to build a reference scale to which localities under investigation will be compared. A detailed example is developed for Early Devonian times. Various plant biocharacters of seven fossiliferous outcrops have been quantified and the subsequent scores of these localities are presented. This example shows that biostratigraphic results obtained by using the quantification method are comparable in precision with dating based on palynological assemblage zones. Evolutionary aspects of the method are also discussed.

Evolutionary history studies depend on having reliable chronologies of macroevolutionary processes. Construction of such chronologies often yields discrepancies between paleontological and molecular dates, which are sometimes viewed as conflicting. Nevertheless, each macroevolutionary process is composed of two main phases: emergence of a trait or clade and success of that trait or clade, which differ in mechanisms, drivers, and types of evidence. Moreover, emergence may be observed as gene divergence (which may be trait-coding or trait-unrelated genes), trait emergence, and clade emergence; whereas success can be observed as increase in abundance, diffusion, and/or diversity or as overall persistence over geologic time. Therefore, to fully and correctly understand any macroevolutionary process, it is of paramount importance to understand what event each date refers to, and how dates of various events and their integration reveal the complexity of macroevolutionary processes. I demonstrate this through three examples: the chronological gap between oxygenic photosynthesis emergence and the Great Oxidation Event, the chronological gap between paleontological and molecular dates of angiosperm emergence, and the evolution of plant silicon accumulation.

Extant species of the genus Equus (e.g., horses, asses, and zebras) have a widespread distribution today on all continents except Antarctica. Extinct species of Equus represented by fossils were likewise widely distributed in the Pliocene and even more so during the Pleistocene. In order to understand the efficacy of “big data” for (paleo)biogeographic analyses, location records (latitude, longitude) and fossil occurrences for the genus Equus were mined and further explored from six databases, including iDigBio, Paleobiology Database, VertNet, BISON, Neotoma, and GBIF. These were chosen from a priori knowledge of where relevant data might be aggregated. We also realized that these databases have different objectives and data sources and therefore would provide a useful comparative study of the widespread taxon Equus in space and time.

The mining of Equus data from these six sources yielded a combined total of 123.8 K location records, including 116.2K fossil specimens. These include individual points that are unique, that is, only occurring in one of these databases, and those that are duplicated in multiple databases. Of the six databases, three (iDigBio, Paleobiology Database, and GBIF) were judged to be the most useful in the Equus use case. Most of the databases are biased toward North American records, thus limiting the reconstruction of the actual distribution of the genus Equus in space and time outside of this continent. Although Equus has a large number of digitally accessible records, fundamentally interesting questions pertaining to evolutionary dynamics and extinction geography are still a challenge for these kinds of biodiversity databases due primarily to the lack of sufficiently dense and precise temporal data.

The fossil record is spatiotemporally heterogeneous: taxon occurrence data have patchy spatial distributions, and this patchiness varies through time. Large-scale quantitative paleobiology studies that fail to account for heterogeneous sampling coverage will generate uninformative inferences at best and confidently draw wrong conclusions at worst. Explicitly spatial methods of standardization are necessary for analyses of large-scale fossil datasets, because nonspatial sample standardization, such as diversity rarefaction, is insufficient to reduce the signal of varying spatial coverage through time or between environments and clades. Spatial standardization should control both geographic area and dispersion (spread) of fossil localities. In addition to standardizing the spatial distribution of data, other factors may be standardized, including environmental heterogeneity or the number of publications or field collecting units that report taxon occurrences. Using a case study of published global Paleobiology Database occurrences, we demonstrate strong signals of sampling; without spatial standardization, these sampling signatures could be misattributed to biological processes. We discuss practical issues of implementing spatial standardization via subsampling and present the new R package divvy to improve the accessibility of spatial analysis. The software provides three spatial subsampling approaches, as well as related tools to quantify spatial coverage. After reviewing the theory, practice, and history of equalizing spatial coverage between data comparison groups, we outline priority areas to improve related data collection, analysis, and reporting practices in paleobiology.