Conodonts were a highly diverse and abundant vertebrate group whose fossils are found in marine Paleozoic and Triassic strata around the world. They inhabited environments ranging from lagoons to open oceans and are represented by a wide variety of dental morphologies. Conodonts may have filled many different ecological niches and represent a significant proportion of nekton before the Devonian. Despite this, very little is known about trophic ecology of conodonts. While morphological diversity suggests a complex trophic structure within conodont communities, there is little evidence to support dietary niche partitioning among conodonts.

We tested the hypothesis that individual conodont taxa occupied different trophic niches, using Sr/Ca and Ba/Ca ratios preserved in the dental elements of assemblages from Silurian strata of Gotland, Sweden. Sr/Ca and Ba/Ca have been shown to vary in vertebrate skeletal tissues depending on trophic positioning, although biological and environmental conditions can affect these ratios. Environmental influences were minimized by examining entire conodont communities from a tropical epeiric sea and by measuring strontium isotope ratios using thermal ionization mass spectrometry in the most metropolitan taxon (Ozarkodina confluens). Composition of white matter, a tissue unique to conodonts, was also analyzed using microprobe analysis, revealing significantly lower Sr concentrations than in surrounding lamellar tissue, suggesting taxon-specific histology should be considered when analyzing conodonts for geochemical data. Excluding taxa with highly variable quantities of white matter, the results show that each taxon preserves different Sr/Ca and Ba/Ca ratios with limited overlap, indicating variation in trophic positioning.

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.

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.

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.

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.

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.

Previous descriptions and interpretations of the Cycadeoidea pollen organ are followed by a new interpretation based upon its presumed neotenous condition. The sexually mature pollen organ represents a juvenile morphological condition that persisted into the adult stage of the organism and was inherited ultimately from a seed fern ancestor. The neotenous organ is considered to have come into existence abruptly and the altered morphology to have been achieved without an intervening series of intermediates. Abrupt establishment of higher taxa by heterochrony constitutes a mechanism capable of decoupling microevolutionary processes from macrevolutionary differences and poses a distinct challenge to current evolutionary theory.

The record of the taxonomic evolution of North American ungulates is critical to our understanding of mammalian evolution and environmental change throughout the Cenozoic. The distribution of sampling in the ungulate fossil record over time and geographic space and the degree to which this biases the observed patterns of taxonomic evolution is poorly understood. To address these issues, I placed fossil collections and occurrences drawn from the Paleobiology Database into 2-Myr time intervals between 55 and 1 Ma. I determined the variation in numbers of fossil collections and occurrences, using three metrics to measure geographic variation: first, the area of the convex hull containing all collections in an interval, to determine the areal coverage of sampling; second, the mean pairwise geographic distance among collections as a measurement of the dispersion of collections within that area; and third, the interval-to-interval migration of the geographic centroid of all collections, to calculate changes in the geographic location of sampling. Each of these showed considerable variation over the Cenozoic, and both the area of the convex hull (ACH) encompassing all collections in an interval, and mean pairwise distance (MPWD) among them showed increasing trends over time.

To minimize the effect of variation in numbers of fossil samples over time, I used standard sample-standardization procedures. To minimize the effect of geographic variation in sampling over time, I standardized the area of sampling among intervals. I also employed both standardizations sequentially. Each standardization procedure had surprisingly little effect on observed patterns of taxonomic richness and rates. This indicates that, for North American ungulates, neither variation in number nor geographic distribution of fossil samples exerts an overwhelming influence on perceived macroevolutionary patterns. These results confirm the ungulate fossil record as a critical and faithful record for our understanding of Cenozoic environmental change and the mammalian evolutionary response.

Recent studies of eurypterid paleoecology suggest that formation of eurypterid Lagerstätten in the mid-Paleozoic of Laurentia was controlled by the presence of an ecological–taphonomic window that recurred predictably in nearshore, marginal environments during transgressions. We tested this hypothesis by performing a high-resolution taxonomic, environmental, and stratigraphic survey and quantitative analysis of all Silurian–Lower Devonian eurypterid-bearing intervals in the Appalachian basin, the most prolific region for eurypterid remains in the world. Canonical correspondence analysis of sedimentological and faunal associations revealed a strong lithologic gradient between groupings of eurypterid genera and associated taxa across the basin, and a significant association of eurypterids with microbialites (thrombolites, stromatolites) and evaporitic structures. Field observations confirmed that, stratigraphically, eurypterids in the basin frequently occur above the microbialite structures and beneath evaporites and other indicators of increased salinity or subaerial exposure. Following interpretation of these features within a sequence stratigraphic framework, we present a preservational model in which (1) eurypterids inhabited nearshore settings following freshening conditions concomitant with minor transgressions, (2) their remains were subsequently buried by storms or microbialite sediment baffling, and (3) subsequent long-term preservation of tissues was facilitated by regression and cyclical shallowing-up successions that promoted hypersalinity and anoxia. In the central and southern region of the basin, where microbial structures and evidence for hypersalinity are less common, a similar pattern of cyclical shallowing-upward deposition within eurypterid-bearing units holds. Thus, eurypterid preservation appears to reflect a combination of ecological preferences and abiotic conditions that promoted inhabitation and eventual preservation within the same setting. This study provides the first quantitative support for a sea level–based control on preservation of eurypterids and adds to the growing body of evidence that suggests that analysis of exceptional preservation in the fossil record benefits from interpretation within a sequence stratigraphic framework.

Beta diversity quantifies the spatial structuring of ecological communities and is a fundamental partition of biodiversity, central to understanding many macroecological phenomena in modern biology and paleobiology. Despite its common application in ecology, studies of beta diversity in the fossil record are relatively limited at regional spatial scales that are important for understanding macroevolutionary processes. The spatial scaling of beta diversity in the fossil record is poorly understood, but has significant implications due to temporal variation in the spatial distribution of fossil collections and the large spatiotemporal scales typically employed. Here we test the spatial scaling of several common measures of beta diversity using the Cenozoic shallow-marine molluscan fossil record of New Zealand and derive a spatially standardized time series of beta diversity. To measure spatial scaling, we use and compare grid-cell occupancy based on an equal-area grid and summed minimum spanning tree length, both based on reconstructed paleocoordinates of fossil collections. We find that beta diversity is spatially dependent at local to regional scales, regardless of the metric or spatial scaling utilized, and that spatial standardization significantly changes apparent temporal trends of beta diversity and, therefore, inferences about processes driving diversity change.

For centuries, paleontologists have sought functional explanations for the uniquely complex internal walls (septa) of ammonoids, extinct shelled cephalopods. Ammonoid septa developed increasingly complex fractal margins, unlike any modern shell morphologies, throughout more than 300 million years of evolution. Some have suggested these morphologies provided increased resistance to shell-crushing predators. We perform the first physical compression experiments on model ammonoid septa using controlled, theoretical morphologies generated by computer-aided design and 3D printing. These biomechanical experiments reveal that increasing complexity of septal margins does not increase compression resistance. Our results raise the question of whether the evolution of septal shape may be tied closely to the placement of the siphuncle foramen (anatomic septal hole). Our tests demonstrate weakness in the centers of uniformly thick septa, supporting work suggesting reinforcement by shell thickening at the center of septa. These experiments highlight the importance of 3D reconstruction using idealized theoretical morphologies that permit the testing of long-held hypotheses of functional evolutionary drivers by recreating extinct morphologies once rendered physically untestable by the fossil record.

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.

The use of sclerobiosis as a tool for paleoenvironmental and paleoecological research is undermined by a lack of comparable methods for sclerobiont data collection and analysis. We present a new method for mapping sclerobiont distributions across any host, and offer an example of how the method may be used to interpret sclerobiont data in relation to host orientation. This approach can also be used to assess the suitability of beds and fossil material for paleoenvironmental reconstruction.

A sample of 150 encrusted dorsibiconvex atrypide brachiopods were selected from six beds in the Waterways Formation (latest Givetian – Early Frasnian; Alberta, Canada). The dorsal and ventral valves of each brachiopod were photographed. Sclerobiont taxa were mapped onto the photographs, and the maps were used to create stacked images with each of the 25 brachiopod specimens from each bed. Based on the life orientation of dorsibiconvex atrypides, three zones were designated on the host: the post mortem zone, (only available to sclerobionts after death and reorientation of the host); the shaded zone (brachial valve, excluding the post mortem zone); and the exposed zone (ventral valve).

Randomization simulation results indicate that all beds likely exhibit non random encrustation patterns, and corroborate the hypotheses that: (1) much of the encrustation occurred while the hosts were alive, and (2) these beds and fossils have experienced little physical reworking or transport and would be suitable for paleoenvironmental analysis. Mapping sclerobionts across hosts can serve as a unifying method to increase the recognition and use of sclerobiosis in paleontological studies.

A basic hypothesis in extinction theory predicts that more abundant taxa have an evolutionary advantage over less abundant taxa, which should manifest as increased survivorship during major extinction events and longer fossil-record durations. Despite this, various paleontologic studies have found conflicting patterns, indicating a more complex relationship between abundance and extinction in the geologic past. This study tests the relationship between abundance and extinction among brachiopod genera within seven third-order depositional sequences spanning the Late Ordovician to Early Silurian (Katian–Aeronian) of the Cincinnati Arch.

Contrary to predictions, abundance is not positively correlated with duration in this study. Abundance and duration range from strongly negatively correlated to uncorrelated depending on the spatial scale of analysis and the geologic intervals included, but correlations never indicate that abundance is an evolutionary advantage. In contrast, abundance was an advantageous trait prior to the Ordovician/Silurian extinction, and brachiopods with higher abundances were more likely to survive the event than less abundant brachiopods. While this result is in keeping with common models of extinction, it has not been observed previously at a mass extinction boundary. This may be further evidence that the Ordovician/Silurian extinction was not accompanied by a shift in the macroevolutionary selectivity regime.

Lunules have evolved independently in several groups of clypeasteroids, including the Rotulidae, Astriclypeidae, Mellitidae and Scutasteridae. In this paper, only the monophyletic assemblage Monophorasteridae plus Mellitidae is considered. Lunules result from modifications of the growth patterns of test plates which bring about changes in relative growth in specific directions. It is unnecessary to postulate resorption of the test. Ambulacral lunules, which are known to have hydrodynamic functions, originated as part of a series of changes: 1) bifurcation of food grooves, 2) formation of pressure drainage channels, 3) lobulation of the ambitus (as in Monophoraster), 4) complete lunule formation. The anal lunule shares the hydrodynamic function but arose separately, as a developmental aberration. The position of the periproct is highly variable but it is most commonly located at the junction of suture lines, at plate corners. In Scutella it is located between the first and second post-basicoronal plates. In Monophoraster the periproct is located further forward, between the first post-basicoronal plates, and the small anal lunule occupies the junction point between first and second post-basicoronals. It is hypothesized that the anal lunule originated as an all-or-nothing event following the forward migration of the periproct and failure to resume normal plate growth at the sutural junction point between first and second post-basicoronal plates. Its walls were derived from paired interambulacral supports. The hypothesis is discussed in connection with the ontogenetic formation of the anal lunule in living mellitid sand dollars.

Species diversity is a product of both biological interactions and physical-geographic factors. A model which eliminates biological interactions as a determinant of diversity can be used to assess the potential impact of physical and geographic factors operating alone on temporal trends in diversity (succession) and on spatial differences along latitudinal or environmental gradients. If a biologically neutral model can generate diversity patterns similar to those we observe, it can be argued that traditional hypotheses which depend on biological interactions are not the most parsimonious explanations for diversity patterns. If the neutral model approach supports a spectrum of possible hypotheses, perhaps the entire concept of species diversity can be discarded as a useful measure of community structure.

An earlier analysis of this type supported the notion that diversity patterns can be adequately explained by physical disturbance, global geography gradients, and stochastic immigration and extinction events. Our neutral model analysis does not affirm this view. We argue that the coarse level at which simple colonization models operate does not permit strong conclusions regarding the underlying mechanisms. Detailed community analyses may provide the best evidence for the nature of structuring processes. Based on the evidence available in the literature, species diversity is likely to be generated by a small number of biological processes interacting with physical-geographic gradients.

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.

The effects of overdispersion and zero inflation (e.g., poor model fits) can result in misinterpretation in studies using count data. These effects have not been evaluated in paleoecological studies of predation and are further complicated by preservational bias and time averaging. We develop a hierarchical Bayesian framework to account for uncertainty from overdispersion and zero inflation in estimates of specimen and predation trace counts. We demonstrate its application using published data on drilling predators and their prey in time-averaged death assemblages from the Great Barrier Reef, Australia.

Our results indicate that estimates of predation frequencies are underestimated when zero inflation is not considered, and this effect is likely compounded by removal of individuals and predation traces via preservational bias. Time averaging likely reduces zero inflation via accumulation of rare taxa and events; however, it increases the uncertainty in comparisons between assemblages by introducing variability in sampling effort. That is, there is an analytical cost with time-averaged count data, manifesting as broader confidence regions. Ecological inferences in paleoecology can be strengthened by accounting for the uncertainty inherent to paleoecological count data and the sampling processes by which they are generated.