The hypothesis that there is a characteristic rate of exponential radiation for the Insecta is tested for forty-nine of the most diverse family-level taxa in the Coleoptera, Diptera, and Hymenoptera. Data used are estimates of Recent species diversity and age of first appearance in the fossil record, as in the original statement of the hypothesis by Stanley (1979). There is no evidence of a characteristic radiation rate for the insects. A more exact method is proposed for identifying clades that are radiating exponentially. When applied to Stanley's data for the mammalian family Bovidae, it indicates that the Bovidae are not radiating exponentially at a characteristic rate. Future research might emphasize the reliable identification of clades that are radiating exponentially, prior to drawing conclusions about rates that might characterize higher taxa.

Lungfishes are known for, and indeed take their name from, their bimodal respiratory abilities. All three extant genera can use their lungs to extract oxygen from the atmosphere, although their reliance upon this capability differs among taxa. Lungs are considered primitive for the Osteichthyes, however the distinctive buccal pump mode of air gulping exhibited by extant lungfishes appears to be a specialization. It is associated with a number of derived skeletal characters (cranial ribs, long parasphenoid stalk, midline gap between palatal tooth plates) that first appeared during the Devonian. These have been described individually, but in no Devonian lungfish has their three-dimensional (3D) spatial relationship been reconstructed and analyzed. Here we present the 3D morphology of Rhinodipterus, a Mid-Late Devonian lungfish from Australia and Europe, based on synchrotron tomography and conventional microtomography scans.

Unlike less crownward contemporaneous lungfishes such as Griphognathus and Chirodipterus, Rhinodipterus has a full set of skeletal buccal pump components that can be directly compared to those of extant lungfishes, suggesting that it made more extensive use of air breathing than other Gogo or Bergisch Gladbach genera. This is interesting in relation to the environmental context as Gogo and Bergisch Gladbach are both marine, contrasting with the frequently hypoxic tropical to subtropical fresh water environments inhabited by modern lungfishes. The evolution of buccal pump-supported lung ventilation was evidently not necessarily associated with a transition to non-marine habitats.

Numerous methods have been developed to estimate the position of a mass extinction boundary while accounting for the incompleteness of the fossil record. Here we describe the point estimator and confidence interval for the extinction that are optimal under the assumption of uniform preservation and recovery potential, and independence among taxa. First, one should pool the data from all taxa into one combined “supersample.” Next, one can then apply methods proposed by Strauss and Sadler (1989) for a single taxon. This gives the optimal point estimator in the sense that it has the smallest variance among all possible unbiased estimators. The corresponding confidence interval is optimal in the sense that it has the shortest average width among all possible intervals that are invariant to measurement scale. These optimality properties hold even among methods that have not yet been discovered. Using simulations, we show that the optimal estimators substantially improve upon the performance of other existing methods. Because the assumptions of uniform recovery and independence among taxa are strong ones, it is important to assess to what extent they are satisfied by the data. We demonstrate the use of probability plots for this purpose. Finally, we use simulations to explore the sensitivity of the optimal point estimator and confidence interval to nonuniformity and lack of independence, and we compare their performance under these conditions with existing methods. We find that nonuniformity strongly biases the point estimators for all methods studied, inflates their standard errors, and degrades the coverage probabilities of confidence intervals. Lack of independence has less effect on the accuracy of point estimates as long as recovery potential is uniform, but it, too, inflates the standard errors and degrades confidence interval coverage probabilities.

Analyses of ancient food webs reveal important paleoecological processes and responses to a range of perturbations throughout Earth's history, such as climate change. These responses can inform our forecasts of future biotic responses to similar perturbations. However, previous analyses of ancient food webs rarely accounted for key differences between modern and ancient community data, particularly selective loss of soft-bodied taxa during fossilization. To consider how fossilization impacts inferences of ancient community structure, we (1) analyzed node-level attributes to identify correlations between ecological roles and fossilization potential and (2) applied selective information loss procedures to food web data for extant systems. We found that selective loss of soft-bodied organisms has predictable effects on the trophic structure of “artificially fossilized” food webs because these organisms occupy unique, consistent food web positions. Fossilized food webs misleadingly appear less stable (i.e., more prone to trophic cascades), with less predation and an overrepresentation of generalist consumers. We also found that ecological differences between soft- and hard-bodied taxa—indicated by distinct positions in modern food webs—are recorded in an early Eocene web, but not in Cambrian webs. This suggests that ecological differences between the groups have existed for ≥48 Myr. Our results indicate that accounting for soft-bodied taxa is vital for accurate depictions of ancient food webs. However, the consistency of information loss trends across the analyzed food webs means it is possible to predict how the selective loss of soft-bodied taxa affects food web metrics, which can permit better modeling of ancient communities.

The work of Arthur J. Boucot (vale, 1924–2017) is foundational to Devonian biogeography. Cited more than 1000 times and mentioned in excess of 105,000 times, his classification of the Old World, Eastern Americas, and Malvinokaffric realms has become the framework for Devonian biogeographic and faunal studies. This study is the first to test and quantify the biotic area classification using the distributional data of trilobite, brachiopod, and fish taxa across the Lower, Middle, and Late Devonian. The biotic similarity analysis of 676 genera (ca. 30,000 occurrences) of late Silurian (Ludlow–Pridoli) and Devonian taxa represents the largest quantification of distributional data undertaken in Devonian biogeography. The temporal area approach (TAAp), applied here for the first time, failed to recover in the area phenogram many of the realms and regions used by Devonian biogeographers, excepting the Malvinokaffric realm, across the entirety of the Devonian. The Old World and Western Gondwana realms are found to be artificial and in need of revision. Without natural areas, bioregionalizations (area classifications) become unstable and unusable, resulting in authors creating further artificial regions and provinces. This study offers a better solution, namely, to identify artificial areas in existing area classifications using TAAp and biotic similarity analysis and revise them. Area classifications are modeled on taxonomic process (Wilson and Brown 1953; Starrett 1958) and the movement toward revision is necessary to break the cycle of reinvention of realms and regions within bioregionalization.

Multivariate methods such as cluster analysis and ordination are basic to paleoecology, but the messy nature of fossil occurrence data often makes it difficult to recover clear patterns. A recently described faunal similarity index based on the Forbes coefficient improves results when its complement is employed as a distance metric. This index involves adding terms to the Forbes equation and ignoring one of the counts it employs (that of species found in neither of the samples under consideration). Analyses of simulated data matrices demonstrate its advantages. These matrices include large and small samples from two partially overlapping species pools. In a cluster analysis, the widely used Dice coefficient and the Euclidean distance metric both create groupings that reflect sample size, the Simpson index suggests large differences that do not exist, and the corrected Forbes index creates groupings based strictly on true faunal overlap. In a principal coordinates analysis (PCoA) the Forbes index almost removes the sample-size signal but other approaches create a second axis strongly dominated by sample size. Meanwhile, species lists of late Pleistocene mammals from the United States capture biogeographic signals that standard ordination methods do recover, but the adjusted Forbes coefficient spaces the points out more sensibly. Finally, when biome-scale lists for living mammals are added to the data set and extinct species are removed, correspondence analysis misleadingly separates out the biome lists, and PCoA based on the Dice coefficient places them to the edge of the cloud of fossil assemblage data points. PCoA based on the Forbes index places them in more reasonable positions. Thus, only the adjusted Forbes index is able to recover true biological patterns. These results suggest that the index may be useful in analyzing not only paleontological data sets but any data set that includes species lists having highly variable lengths.

Many apparent homeomorphs have been recognized among genera assigned to the Paleozoic stenolaemate bryozoan orders Trepostomata and Cryptostomata. Morphologies are evaluated to determine whether such homeomorphy should be expected, or is likely to have resulted from incomplete taxonomic analysis. Historical and constructional constraints prevalent in the phylum appear to provide ample opportunity for adaptive convergence, a conclusion supported by the occurrence of similar morphologies in distantly related post-Paleozoic bryozoans.

Cryptostomes are distinguished in part by the presence of restricted budding loci. Loci pattern is hypothesized to be the key innovation in the establishment of the clade and the pattern also is considered critical to the evolution of the relatively slender branches and short zooecia typical of cryptostomes.

Loci development does not provide a taxonomic panacea. This is because the hypothesis does not preclude convergent evolution of broadly similar restricted loci in Paleozoic non-cryptostome groups, although presumably the cryptostomes in general would have been best equipped to succeed under conditions favoring restricted loci.

Environmental conditions for the cryptostomes in general and changing life history strategies for rhabdomesoids in particular provide possible controlling factors for the evolution of these bryozoans.

The stratophenetic approach to phylogeny reconstruction, developed in numerous recent papers by Gingerich, has its strongest data base in comprehensive collections from early Eocene rocks in the Big Horn Basin of northwestern Wyoming. Other examples have been criticised for lack of well-documented collection data, making suitable testing of stratophenetic interpretations of evolutionary modes virtually impossible.

Study of Hyopsodus from the middle Eocene Bridger Formation of southwestern Wyoming provides new collection data. Adequate stratigraphic documentation is available, so Gingerich's graphical presentation can be essentially duplicated. In contrast to the complex branching pattern Gingerich interpreted from his data, Bridger Formation Hyopsodus data seems to show little size change through approximately one million years. This stasis or equilibrium condition, which is considered rare by Bookstein et al. (1978), is the only well developed pattern in Bridger Hyopsodus.

As the climate changes and ecosystems shift toward novel combinations of species, the methods and metrics of conservation science are becoming less species-centric. To meet this growing need, marine conservation paleobiologists stand to benefit from the addition of new, taxon-free benthic indices to the live–dead analysis tool kit. These indices, which were developed to provide actionable, policy-specific data, can be applied to the readily preservable component of benthic communities (e.g., mollusks) to assess the ecological quality status of the entire community. Because these indices are taxon-free, they remain applicable even as the climate changes and novel communities develop—making them a potentially valuable complement to traditionally applied approaches for live–dead analysis, which tend to focus on maintaining specific combinations of species under relatively stable environmental conditions. Integrating geohistorical data with these established indices has potential to increase the salience of the live–dead approach in the eyes of resource managers and other stakeholders.

Geographic distribution, species duration, and population size of 41 bivalve species which coexisted during the uppermost biozone (Sciponoceras gracile zone) of the Cenomanian were measured. Average species duration and correlation between geographic distribution, species duration, and population size were computed for five groups: total, infaunal, epifaunal, cemented epifaunal, and uncemented epifaunal bivalves.

Average species duration for all bivalves is less than 2 Myr. Infaunal bivalves averaged slightly higher, epifaunal slightly lower. Positive correlation was measured between species duration and geographic distribution for all five groups and statistical significance of the relationships was demonstrated for all but infaunal forms. In general no correlation was found between species duration and population size, and between population size and geographic distribution with the following exceptions: species duration and population size were significantly correlated for infaunal bivalves and uncemented epifaunal bivalves, and population size and geographic distribution were correlated for uncemented bivalves. Because these 41 fossil species coexisted at the same time and were distributed in the same space, differences in the relationships of species duration, geographic distribution and population among the various life habits indicate differences in ecological strategy and the evolutionary consequences of that difference.

The field of sclerochronology has long been known to paleobiologists. Yet, despite the central role of growth rate, age, and body size in questions related to macroevolution and evolutionary ecology, these types of studies and the data they produce have received only episodic attention from paleobiologists since the field's inception in the 1960s. It is time to reconsider their potential. Not only can sclerochronological data help to address long-standing questions in paleobiology, but they can also bring to light new questions that would otherwise have been impossible to address. For example, growth rate and life-span data, the very data afforded by chronological growth increments, are essential to answer questions related not only to heterochrony and hence evolutionary mechanisms, but also to body size and organism energetics across the Phanerozoic. While numerous fossil organisms have accretionary skeletons, bivalves offer perhaps one of the most tangible and intriguing pathways forward, because they exhibit clear, typically annual, growth increments and they include some of the longest-lived, non-colonial animals on the planet. In addition to their longevity, modern bivalves also show a latitudinal gradient of increasing life span and decreasing growth rate with latitude that might be related to the latitudinal diversity gradient. Is this a recently developed phenomenon or has it characterized much of the group's history? When and how did extreme longevity evolve in the Bivalvia? What insights can the growth increments of fossil bivalves provide about hypotheses for energetics through time? In spite of the relative ease with which the tools of sclerochronology can be applied to these questions, paleobiologists have been slow to adopt sclerochronological approaches. Here, we lay out an argument and the methods for a path forward in paleobiology that uses sclerochronology to answer some of our most pressing questions.

An increasing number of unexpectedly diverse benthic communities are being reported from microbially precipitated carbonate facies in shallow-marine platform settings after the end-Permian mass extinction. Ostracoda, which was one of the most diverse and abundant metazoan groups during this interval, recorded its greatest diversity and abundance associated with these facies. Previous studies, however, focused mainly on taxonomic diversity and, therefore, left room for discussion of paleoecological significance. Here, we apply a morphometric method (semilandmarks) to investigate morphological variance through time to better understand the ecological consequences of the end-Permian mass extinction and to examine the hypothesis that microbial mats played a key role in ostracod survival. Our results show that taxonomic diversity and morphological disparity were decoupled during the end-Permian extinction and that morphological disparity declined rapidly at the onset of the end-Permian extinction, even though the high diversity of ostracods initially survived in some places. The decoupled changes in taxonomic diversity and morphological disparity suggest that the latter is a more robust proxy for understanding the ecological impact of the extinction event, and the low morphological disparity of ostracod faunas is a consequence of sustained environmental stress or a delayed post-Permian radiation. Furthermore, the similar morphological disparity of ostracods between microbialite and non-microbialite facies indicates that microbial mats most likely represent a taphonomic window rather than a biological refuge during the end-Permian extinction interval.

Chitinous arm hooks (onychites) of belemnoid coleoid cephalopods are widely distributed in Mesozoic sediments. Due to their relative abundance and variable morphology compared with the single, bullet-shaped, belemnite rostrum, arm hooks came into the focus of micropaleontologists as a promising index fossil group for the Jurassic–Cretaceous rock record and have been the target of functional, ecological, and phylogenetic interpretations in the past. Based on three well-preserved arm crowns of the Toarcian diplobelid Chondroteuthis wunnenbergi, we analyzed the shape of a total of 87 micro-hooks. The arm crown of Chondroteuthis is unique in having uniserial rather than biserial hooks. The first application of elliptic Fourier shape analysis to the arm weapons of belemnoid coleoids allows for the distinction of four micro-hook morphotypes and the quantification of shape variation within these morphotypes. Based on the best-preserved arm crown, we reconstructed the distribution of morphotypes within the arm crown and along a single arm. Our quantitative data support former observations that smaller hooks were found close to the mouth and at the most distal arm parts, while the largest hooks were found in the central part of the arm crown. Furthermore, we found a distinct arm differentiation, as not every arm was equipped with the same hook morphotype. Here, we report the functional specialization of the belemnoid arm crown for the first time and speculate about the potential function of the four morphotypes based on comparisons with modern cephalopods. Our analyses suggest a highly adapted functional morphology and intra-individual distribution of belemnoid hooks serving distinct purposes mainly during prey capture.

The tailbend, a downward flexure of the vertebral column, is a prominent feature of post-Triassic ichthyosaurs. However, due to the vagaries of preservation and of preparation, its presence cannot be verified in all skeletons. Computed tomography offers a solution to the problem by identifying the wedge-shaped centra that contribute to the tailbend. Leptopterygius tenuirostris appears to have a tailbend, albeit slightly downturned. There is also evidence that its close relative Eurhinosaurus longirostris had a tailbend, too.

Hadrosaurid dinosaurs, the dominant large-bodied terrestrial herbivores in most Laurasian Late Cretaceous ecosystems, have an exceptional fossil record consisting of many species known from partial ontogenetic series, making them an ideal clade with which to conduct life-history studies. Previous research considered the Dinosaur Park Formation (DPF) of Alberta as an attritional, or time-averaged, sample and interpreted size–frequency distribution of long bones collected from the DPF with three size classes to suggest that hadrosaurids from the DPF attained near-asymptotic body size in under 3 years. This conflicted with previously published osteohistological estimates of 6+ years for penecontemporaneous hadrosaurids from the Two Medicine Formation (TMF) of Montana, suggesting either extreme variation in hadrosaurid growth rates or that size–frequency distributions and/or osteohistology and growth modeling inaccurately estimate ontogenetic age.

We tested the validity of the previously proposed size–age relationship of hadrosaurids from the DPF by significantly increasing sample size and combining data from size–frequency distributions and osteohistology across multiple long-bone elements. The newly constructed size–frequency distributions typically reveal four relatively distinct size–frequency peaks that, when integrated with the osteohistological data, aligned with growth marks. The yearling size class was heavily underrepresented in the size–frequency distribution. If not due to preservation, this suggests that either juvenile (<2 years of age) hadrosaurids from the DPF had increased survivorship following an initially high nestling mortality rate or that yearlings were segregated from adults. A growth-curve analysis revealed asymptotic body size was attained in approximately 7 years, which is consistent with hadrosaurids from the TMF. The data suggest size–frequency distributions of attritional samples underestimate age and overestimate growth rates, but when paired with osteohistology can provide unique life-history insights.

Paleobiological and paleoecological interpretations rely on constraining the temporal resolution of the fossil record. The taphonomic clock, that is, a correlation between the alteration of skeletal material and its age, is an approach for quantifying time-averaging scales. We test the taphonomic clock hypothesis for marine demersal and pelagic fish otoliths from a 10–40 m depth transect on the Mediterranean siliciclastic Israeli shelf by radiocarbon dating and taphonomic scoring. Otolith ages span the last ~8000 yr, with considerable variation in median and range along the transect. Severely altered otoliths, contrary to pristine otoliths, are likely to be older than 1000 yr. For pelagic fish otoliths, at 30 m depth, taphonomic degradation correlates positively with postmortem age. In contrast, no correlation occurs for demersal fishes at 10 and 30 m depth, mostly because of the paucity of very young pristine (<150 yr) otoliths, possibly due to a drop in production over the last few centuries. Contrary to molluscan and brachiopod shells, young otoliths at these depths are little affected and do not show a broad spectrum of taphonomic damage, because those that derive from predation are excreted in calcium- and phosphate-rich feces forming an insoluble crystallic matrix that increases their preservation potential. At 40 m depth, all dated otoliths are very young but rather damaged because of locally chemically aggressive sediments, thus showing no correlation between taphonomic grade and postmortem age. Our results show that local conditions and the target species population dynamics must be considered when testing the taphonomic clock hypothesis.

The fossil record of mammal dentition provides crucial insight into key ecological and functional transitions throughout mammalian evolutionary history. For cetaceans, both extant clades differ markedly from their stem ancestors; neither retains the differentiated dentition or the tribosphenic molars characteristic of Mammalia. We used quantitative measures of dental complexity across fossil and living cetaceans to identify a trend toward dental simplicity through the Neogene. Both extant cetacean clades depart from the ancestral mammalian condition and concurrently converge upon a reduced and simplified dentition; modern mysticetes all have become entirely edentulous (at birth), and living odontocetes possess teeth as single-rooted, conical pegs. These two parallel trends accompany major shifts in feeding strategy (i.e., filter feeding in mysticetes and echolocation in odontocetes), suggesting that these evolutionary innovations for prey acquisition are enabling factors for the loss of prey processing and subsequent convergence on dental simplification.

More than 500 undistorted, unfragmented pedicle valves of Ambocoelia umbonata (Conrad) (Spiriferida, Brachiopoda) were recovered from each of four levels within a Middle Devonian fossil cluster. The fossil cluster, an ellipsoidal shell accumulation measuring one meter in diameter and 2 cm in thickness, was exhumed from an exposure of the Ludlowville Shale (Hamilton Group) of western New York. Size frequency histograms indicate that the brachiopod experienced very high levels of juvenile mortality, due, probably, to the effects of high bottom turbidity. Sedimentological and paleontological evidence indicates that the cluster represents a sequence of in situ benthic associations.

Size-independent variation was estimated by calculating the eigenvalue of the minor axis on a log-transformed plot of pedicle valve lengths and widths. The eigenvalue technique eliminates the effect of allometrically induced shape changes and is applicable to multicharacter analyses of morphological variability.

Size-independent variability among the larger individuals of Ambocoelia decreases in successively younger cluster populations. The decrease is not correlated with any observed or inferred change in substratum, diversity, equitability or turbidity.

Recovery of trilobite faunas following a mass extinction during the Upper Cambrian Sunwaptan Stage (equal to the “Ptychaspid Biomere”) involved increases in both within-habitat (alpha) diversity and in levels of biofacies differentiation. This pattern parallels, at a lower hierarchical level, the global increase in Phanerozoic marine invertebrate species richness (Sepkoski et al. 1981), which was influenced by changes in provinciality and in alpha diversity. Diversification slowed markedly in subtidal shelf carbonate habitats in the upper half of the stage, but continued in carbonate shelf margin environments, possibly due to higher rates of immigration from off-shelf sites. Trilobite clades appearing early in the stage have low centers of gravity, reflecting high immigration rates during and shortly after an interval of mass extinction. Expansion of shallow-water carbonate deposition in the upper half of the stage was accompanied by the spread of carbonate bank biofacies. Families occurring in these biofacies gradually increased in species richness and tended to have high centers of gravity.

With a single complete mandible and 56 mandibular symphyseal fragments of various sizes, the Late Cretaceous Hungarian azhdarchid material has been considered one of the most extensive monospecific pterosaur assemblages in the world. Representing a broad size range, these elements have been thought to demonstrate a developmental series of Bakonydraco galaczi. As such, they were ideal to test whether absolute size and/or morphology reliably indicate relative ontogenetic stages in this pterosaur. Forty-five specimens were selected for multivariate morphometrics and classified into four size classes. After acquiring the morphometric data set, we thin-sectioned eight symphyses representing all size groups and classified them into relative ontogenetic stages based on qualitative microstructural inspection prior to quantitative histological analyses. Microstructural characters suggestive of developmental state were then quantified for intra- and interindividual uni- and multivariate analyses to test the correspondence among the results of qualitative and quantitative analyses. In contrast to our expectations, histological features identified the smallest specimen as an adult and not an early juvenile. The substantial size difference between this specimen and other adults, along with its distinct microanatomical and histological features, implies the presence of at least two pterosaur taxa in this symphysis assemblage. This hypothesis is further supported by multivariate morphometrics, which separate the smallest symphyses from all other specimens that form one continuous group. Although the latter group also shows considerable size variability in corresponding ontogenetic stages, this suggests developmental plasticity rather than the presence of even more taxa, and indicates that symphysis size and morphology are poor indicators of skeletal maturity in these animals. Hence, bone histology is an important independent test of the assessment of ontogenetic stage using size and morphology.