Detailed analysis of the stratigraphic ranges of Devonian rugose coral genera within the Old World and Eastern Americas Realms gives new information on faunal extinctions and other bioevents in both realms. Various origination and extinction metrics are calculated from tabulations of occurrences in each stage. The most significant faunal changes were near or at the ends of the Lochkovian and Frasnian stages. The former marks the gradual transition from dominance by Silurian families and genera to the characteristic Devonian coral assemblages; the latter marks the virtual extinction of the Devonian families and genera. Other coral events are related to the two major changes.

The data provide new bases for comparing the histories of the two realms. Most of the events are recorded in both, giving support to previous suggestions that the causes were worldwide. The coral record shows an increase (probably episodic) in environmental deterioration persisting through the Middle Devonian and culminating in extinction at the end of the Frasnian. Eustatic sea level fluctuations may have caused the precursor events and a bolide impact may have caused the end-Frasnian extinction.

Geologically rapid climate change is anticipated to increase extinction risk nonuniformly across the Earth's surface. Tropical species may be more vulnerable than temperate species to current climate warming because of high tropical climate velocities and reduced seawater oxygen levels. To test whether rapid warming indeed preferentially increased the extinction risk of tropical fossil taxa, we combine a robust statistical assessment of latitudinal extinction selectivity (LES) with the dominant views on climate change occurring at ancient extinction crises. Using a global data set of marine fossil occurrences, we assess extinction rates for tropical and temperate genera, applying log ratios to assess effect size and Akaike weights for model support. Among the classical “big five” mass extinction episodes, the end-Permian mass extinction exhibits temperate preference of extinctions, whereas the Late Devonian and end-Triassic selectively hit tropical genera. Simple links between the inferred direction of climate change and LES are idiosyncratic, both during crisis and background intervals. More complex models, including sampling patterns and changes in the latitudinal distribution of continental shelf area, show tropical LES to be generally associated with raised tropical heat and temperate LES with global cold temperatures. With implications for the future, our paper demonstrates the consistency of high tropical temperatures, habitat loss, and the capacity of both to interact in generating geographic patterns in extinctions.

A survey of the primary xylem anatomy and dimensions of axes (=“stems”) of Upper Devonian plant petrifaction/compression fossils reveals the following: among genera, the primary xylem strand (1) increasingly deviates from a cylindrical geometry (=haplostele) as both the axis diameter and the nonxylem (“ground”) tissue volume increase and (2) increases in geologically younger specimens. (3) The perimeter length to area ratio of the primary xylem strand (or the primary xylem surface area to volume ratio) is dependent upon axis diameter or the area of the ground tissue. This ratio (4) decreases in evolutionary lineages that involve the appearance of larger plants that maintain a haplostelar anatomy (e.g., the evolution of trimerophytes from a rhyniophyte ancestor) or increases in evolutionary lineages involving the appearance of nonhaplostele anatomy (e.g., the evolution of progymnosperms and pteridophytes from trimerophyte ancestors) and (5) the relative amount of primary xylem to ground tissue increases in stratigraphically younger specimens. These trends are interpreted within the context of (1) correlations between the diameter of axes and the complexity of the primary xylem anatomy reported for some extant pteridophytes and (2) changes associated with the evolution of new primary xylem configurations and organographic relationships. Based on the appearance of new ontogenetic patterns and concomitant changes in the anatomy of the primary xylem strand associated with the evolution of derived plant groups, a stria extrapolation of a size to complexity correlation based on ontogenetic studies to the phyletic level is considered suspect.

Reduction of body size is a common response of organisms to environmental stress. Studying the early Toarcian succession in the Lusitanian Basin of Portugal, we tested whether the shell size of benthic marine communities of bivalves and brachiopods changed at and before the global, warming–related Toarcian oceanic anoxic event (T-OAE). Statistical analyses of shell size over time show that the mean shell size of communities decreased significantly before the T-OAE. This trend is distinct in brachiopods and is caused by larger-sized species becoming less abundant over time, whereas it is not significant in bivalves, suggesting a decoupled response to environmental stress. Reductions in shell size precede the decline in standardized sample-level species richness associated with the early Toarcian extinction event. Such decreases in the shell size of marine invertebrates, well before the onset of biodiversity change, suggest that reductions in body size more generally may be a precursor of a subsequent loss of species and turnover at the community level caused by climate change. Sedimentological evidence is against hypoxia as a driver of extinction and the preceding size decrease in the brachiopod fauna in the studied succession, although low oxygen levels are widely held responsible for elevated early Toarcian extinction rates globally. Reduction of mean shell size in brachiopods but stasis in bivalves is difficult to explain with ocean acidification, because experimental work shows that brachiopods can be resilient to lowered pH, albeit long-term metabolic costs and potential evolutionary adaptations are unknown. Rising early Toarcian temperatures in the Lusitanian Basin seem to be a plausible factor in both diversity decline associated with the T-OAE and the preceding reductions in mean shell size, because thermal tolerances in modern bivalves are among the highest within marine invertebrates.

There is a well-established discrepancy between paleontological and molecular data regarding the timing of the origin and diversification of placental mammals. Molecular estimates place interordinal diversification dates in the Cretaceous, while no unambiguous crown placental fossils have been found prior to the end-Cretaceous mass extinction. Here, the completeness of the eutherian fossil record through geological time is evaluated to assess the suggestion that a poor fossil record is largely responsible for the difference in estimates of placental origins. The completeness of fossil specimens was measured using the character completeness metric, which quantifies the completeness of fossil taxa as the percentage of phylogenetic characters available to be scored for any given taxon. Our data set comprised 33 published cladistic matrices representing 445 genera, of which 333 were coded at the species level.

There was no significant difference in eutherian completeness across the Cretaceous/Paleogene (K/Pg) boundary. This suggests that the lack of placental mammal fossils in the Cretaceous is not due to a poor fossil record but more likely represents a genuine absence of placental mammals in the Cretaceous. This result supports the “explosive model” of early placental evolution, whereby placental mammals originated around the time of the K/Pg boundary and diversified soon after.

No correlation was found between the completeness pattern observed in this study and those of previous completeness studies on birds and sauropodomorph dinosaurs, suggesting that different factors affect the preservation of these groups. No correlations were found with various isotope proxy measures, but Akaike information criterion analysis found that eutherian character completeness metric scores were best explained by models involving the marine-carbonate strontium-isotope ratios (87Sr/86Sr), suggesting that tectonic activity might play a role in controlling the completeness of the eutherian fossil record.

Physical models of graptolites have been constructed for a range of morphologies, with emphasis on planar, multiramous forms. The models are life size and have the density of a living graptolite, based on the now-established collagenous nature of the periderm and unavoidable assumptions about the amount of extrathecal tissue present in the living colony. These models have been used to test the two main hypotheses of graptolite life habit developed by Bulman, Rickards, Kirk, and others. Testing of graptoloid models in water suggests that many rhabdosome shapes were designed for passive rotation within the water column. This is caused in the models by a variety of modifications, including changes in thecal and stipe orientation, alterations of colony shape and the addition of vanes and hooks. Rotation would only have been useful when the rhabdosome was in directional motion and the frequency of such modifications seems anomalous if no such movement occurred. Thus movement by some means is required, either passively, by changes in buoyancy, or by automobility. Spiralling action would increase the harvesting path of an individual living on a planar, multiramous colony, making this a theoretically advantageous mode of life for the morphology. It would prevent the individual zooids of scandent biserial and uniserial colonies from feeding from the same narrow band of water.

The marine faunas of tropical America underwent substantial evolutionary turnover in the past 3 to 4 million years in response to changing environmental conditions associated with the rise of the Isthmus of Panama, but the ecological signature of changes within major clades is still poorly understood. Here we analyze the paleoecology of faunal turnover within the family Pectinidae (scallops) over the past 12 Myr. The fossil record for the southwest Caribbean (SWC) is remarkably complete over this interval. Diversity increased from a low of 12 species ca. 10–9 Ma to a maximum of 38 species between 4 and 3 Ma and then declined to 22 species today. In contrast, there are large gaps in the record from the tropical eastern Pacific (TEP) and diversity remained low throughout the past 10 Myr. Both origination and extinction rates in the SWC peaked between 4 and 3 Ma, and remained high until 2–1 Ma, resulting in a 95% species level turnover between 3.5 and 2 Ma. The TEP record was too incomplete for meaningful estimates of origination and extinction rates. All living species within the SWC originated within the last 4 Myr, as evidenced by a sudden jump in Lyellian percentages per faunule from nearly zero up to 100% during this same interval. However, faunules with Lyellian percentages near zero occurred until 1.8 Ma, so that geographic distributions were extraordinarily heterogeneous until final extinction occurred. There were also striking differences in comparative diversity and abundance among major ecological groups of scallops. Free-swimming scallops constituted the most diverse guild throughout most of the last 10 Myr in the SWC, and were always moderately to very abundant. Leptopecten and Argopecten were also highly diverse throughout the late Miocene and early Pliocene, but declined to very few species thereafter. In contrast, byssally attaching scallops gradually increased in both diversity and abundance since their first appearance in our samples from 8–9 Ma and are the most diverse group today. Evolutionary turnover of scallops in the SWC was correlated with strong ecological reorganization of benthic communities that occurred in response to declining productivity and increased development of corals reefs.

Symbiotic relationships involving physical contact between worms and solitary rugosan polyps are recorded by the following structures in North American Late Ordovician corals: (1) Trypanites borings enclosed within septal swellings in two specimens, (2) vermiform grooves and openings along the external wall of one corallum, and (3) a chamber containing a unique brown tube within one individual. These features are indicative, respectively, of commensal boring polychaete annelids that penetrated through coralla, commensal epizoic worms of unknown taxonomic affinity that attached to the side of a polyp, and a tubicolous worm (possibly a polychaete) that was likely a parasitic endozoan. Symbionts comparable to the latter two types are also known from two specimens of Devonian solitary rugose corals.

Indirect evidence suggests that symbioses between solitary rugosans and the worms that produced Trypanites borings as dwelling structures in the sides of coralla were relatively common. However, direct evidence that the hosts were alive has been found in only two corals. In both cases, worms bored through septa within the calices and came into contact with basal surfaces of the polyps, which secreted skeletal material that sealed off the intruders. The rarity of such structures suggests that the encounters were inadvertent. If boring worms favored upcurrent portions of objects in order to maximize feeding benefits and avoid sedimentation, their locations indicate that the concave sides of curved coralla faced toward prevailing currents when in life positions.

“Opportunistic” worms are known to have attached to the sides of polyps only in rare instances when the hosts became temporarily exposed as a result of accidents or abnormalities. This indicates that coralla normally served to shield polyps from colonization by nonboring epizoans.

Worms that apparently extended up through openings in basal surfaces of polyps likely obtained sustenance parasitically within the central cavities. They could have entered the hosts through their mouths, or via the calices when parts of the polyps detached from their coralla and contracted radially. The rarity of this type of relationship in solitary Rugosa suggests that the worms entered inadvertently.

Symbioses involving physical contact between worms and polyps seem to have been rare throughout the history of solitary rugose corals. Both groups apparently tolerated such associations when they did occur, although the rugosans secreted structures in their coralla that served to isolate the symbionts. In doing so, they recorded the presence of worms not likely to be preserved as body fossils. The interpretation of such features provides information on the physiology and ethology of both organisms, on the history of symbiotic relationships, and on the diversity of soft-bodied organisms in ancient environments.

I describe a mathematical model of shells that incorporates simple morphometric measurements, which can be obtained easily from sectioned specimens. The model reconstructs form (size and shape) by describing the path in space traversed by, and the changes in dimensions of, a shell aperture during accretionary growth. The model contains parameters that independently affect components of these two aspects of shell form in directions radially away from (or toward) and along a coiling axis. I also describe a computer program, CerioShell, that uses the mathematical model to graphically simulate shell forms. The program permits the viewing of images from different three-dimensional orientations, using various graphical representations, and provides dimensions of the images it generates. Together, the model and the program allow one to investigate the effects that changes in modes of growth would have on shell form.

Writers as disparate in professional interest as those describing speciation in Drosophila, predicting weather, or supporting creationism have all recently focused on the “sudden” appearance of taxa in the geologic record. In the process, each has cited the punctuated equilibrium argument. Owing to the enormously wide attention which this issue has received since its introduction almost a decade ago, it seems worthwhile to try to place the paleontological and biological evidence in a 1981 perspective. The focus I have chosen is to try to analyze the issue, much as in recent columns in Current Happenings on competitive exclusion, and on larval dispersal and biogeography, in which are put forth alternative explanations for a set of data on a general problem.

Poecilozonites (Gastrelasmus) is an important component of the endemic land snail fauna of Pleistocene Bermuda. The type species P. circumfirmatus Redfield usually occurs in sympatry with its congener P. discrepans Pfeiffer, though each species is found alone at several localities. The species are less alike morphologically where they occur together than where they are allopatric. This allopatric convergence and sympatric divergence strongly suggests the biological interaction known as character displacement, often documented for living populations. The relatively complete fossil record of Bermuda offers advantages for studying this phenomenon. Collections can be made from a variety of microhabitats occupied through time. Statistical analysis of 1,600 individuals collected from more than 100 localities indicates that interspecific variation is primarily a function of the presence or absence of a congener and depends to a lesser degree on microhabitat. P. circumfirmatus undergoes a smaller morphological change between allopatry and sympatry than does P. discrepans. Study of relative abundances suggests that P. discrepans may have been competitively inferior, though no functional bases for differences between the species are known.

Minimum Sample Richness (MSR) is defined as the smallest number of taxa that must be recorded in a sample to achieve a given level of inter-assemblage classification accuracy. MSR is calculated from known or estimated richness and taxonomic similarity. Here we test MSR for strengths and weaknesses by using 167 published mammalian local faunas from the Paleogene and early Neogene of the Quercy and Limagne area (Massif Central, southwestern France), and then apply MSR to 84 Oligo-Miocene faunas from Riversleigh, northwestern Queensland, Australia. In many cases, MSR is able to detect the assemblages in the data set that are potentially too incomplete to be used in a similarity-based comparative taxonomic analysis. The results show that the use of MSR significantly improves the quality of the clustering of fossil assemblages. We conclude that this method can screen sample assemblages that are not representative of their underlying original living communities. Ultimately, it can be used to identify which assemblages require further sampling before being included in a comparative analysis.

Experiments were conducted to test the hypothesis that biomechanical constraints determine the morphology of regular echinoids. Hard-bottom-dwelling Tripneustes gratilla elatensis were transferred to an artifical sandy habitat to evaluate whether the change in substrate affects their height to diameter ratio (H/D). Within 1–2 months their H/D ratio increased significantly. This change was shown to be reversible to some extent. Surgical damage to the ambulacral system of one ray caused inactivation of tubefeet and atrophy of injured ambulacra. Test shape was also affected: the damaged ray was lower, and the nondamaged ambulacra deflected toward the treated one, producing bilateral symmetry as in recorded cases of teratology. Study of T. g. elatensis tetramers showed that while “perfect” tetramery was apparently associated with genetic aberration, “imperfect” tetramery results from mechanical injury at an early ontogenetic stage. Micromorphological study shows that in the longitudinal sutures, normally under tension, long and slender trabeculae develop, associated with long and well-aligned collagenous sutural fibers, while the latitudinal trabeculae and fibers are short and less organized. A mechanical effect is suggested by the oval cross-section of the fiber-anchoring trabeculae. Further, echinoid plates interact like soap bubbles, whereas the entire test behaves like a balloon, fastened to the substrate by the ambulacral tubefeet. All these observations support earlier hypotheses on the biomechanical control of echinoid test growth. A model is proposed in which the expansion of the inner mass, counteracted by the mechanical activity of the ambulacral tubefeet, mesenterial threads, and lantern muscles, affects sutural growth, thus controlling echinoid morphogenesis.

A morphometric survey among regular echinoids reveals an inverse relationship between ambulacral width and relative ambital height. Although both increase of ambulacral width and lowering of ambitus-line are evolutionary trends, it is suggested that they are a response to a mechanical effect. H/D ratio was not related to ambulacral width on the phylogenetic level. It is therefore suggested that the latter correlation is ontogenetically controlled. Aspects of irregular echonoid evolution, such as bilateral symmetry, flattening, and formation of the ambulacral petaloid, also are explained by this model.

We use Fourier analysis and related techniques to investigate the question of periodicities in fossil biodiversity. These techniques are able to identify cycles superimposed on the long-term trends of the Phanerozoic. We review prior results and analyze data previously reduced and published. Joint time-series analysis of various reductions of the Sepkoski Data, Paleobiology Database, and Fossil Record 2 indicate the same periodicity in biodiversity of marine animals at 62 Myr. We have not found this periodicity in the terrestrial fossil record. We have found that the signal strength decreases with time because of the accumulation of apparently “resistant” long-lived genera. The existence of a 62-Myr periodicity despite very different treatment of systematic error, particularly sampling-strength biases, in all three major databases strongly argues for its reality in the fossil record.

A marked retardation of dental ontogeny characterizes the family Elephantidae. As a consequence of this retardation, elephant teeth are subject to the forces of mastication, eruption, and progression while still in a developing and pliant stage. As specimens described here illustrate, the mechanical forces are often sufficient to deform the gross morphology of dentitions. Morphological variation in elephant teeth can be regarded as “fabricational noise”—revealing information about the dynamic context in which the teeth develop. Accordingly, dental variation is less species-specific in elephants than in other mammals. The fossil record may comprise fewer species of elephants than is generally believed, and trends inferred to reflect rapid evolution within this family may in fact reflect phenotypic plasticity.

A general model of species dynamics must incorporate the effects of species number on the processes of speciation and extinction. Previous models make specific assumptions about these effects, but do not consider the effects of dynamics of lower level entities on speciation and extinction rates. A hierarchical model is developed which explicitly describes the effects of energy use by species on speciation and extinction rates. The effects of energy use are represented by parameters that characterize the average effects of energy use by each species in the biota on speciation and extinction rates. The dynamics of the model describe a sigmoidal increase in species number over time, as does the logistic model of species dynamics. However, the mechanisms of those dynamics are assumed to be different in the two models. Empirical analysis of a data set on the diversification of fossil Miocene horses suggests that the logistic and hierarchical models have similar descriptive power. The hierarchical model incorporates insights from recent considerations of the nature of a hierarchical theory of biology. Further progress in developing such a theory will depend on the success with which relationships among levels in the biological hierarchy are able to be defined.

Astogenetic trajectories have constrained evolutionary changes in bryozoans. Rates and timing of astogenetic differentiation have been modified for characters defining the morphology of zooids, subcolonies, and colonies. This paper catalogs 46 examples of bryozoan heterochrony, representing all five skeletonized orders. Heterochrony is inferred to have been a pervasive phenomenon in the evolution of Paleozoic stenolaemates, illustrated by 40 examples, 19 of which produced paedomorphosis and 21 peramorphosis. As a consequence, a restricted range of morphologic states has reappeared repetitively as homeomorphies and evolutionary reversals. Large-scale patterns developed across both geologic time and geographic space reflect variation in heterochronic products irrespective of the developmental processes by which they were achieved. Available evidence indicates that smaller, paedomorphic, and more plastic species inhabited onshore, low-diversity areas. Nonheritable plasticity is inferred to be a correlate of early growth stages and paedomorphosis. Taller, generally peramorphic species with damped plasticity are found in higher diversity, offshore regions. Seven key innovations, which first appeared during the early diversification of bryozoan clades, are peramorphic, and recapitulation was a predominant pattern during their Ordovician radiation. Trends in later phylogeny, on the other hand, have favored paedomorphic derived morphologies, as illustrated by 19 of the 32 examples. Recurrent reverse recapitulation suggests that offshore ancestors frequently gave rise to onshore paedomorphs.

Computer models of the tail of Apatosaurus louisae show it could reach supersonic velocities, producing a noise analogous to the “crack” of a bullwhip. Similarity in tail structure suggests this was feasible for other diplodocids, and possibly for unrelated sauropods like Mamenchisaurus and the dicraeosaurids. Lengthening of caudal vertebrae centra between positions 18 and 25 is consistent with adaptation to the stresses generated by such tail motion, as is coossification of vertebrae via diffuse idiopathic skeletal hyperostosis (DISH), which occurs in the same region in about half the specimens. The noise produced may have been used for defense, communication, intraspecific rivalry, or courtship, in which case supersonic “cracking” may have been a sexually dimorphic feature. Comparisons with the club-bearing tails of the sauropods Shunosaurus lii and Omeisaurus tianfuensis show the diplodocid whiplash tail was not well adapted as a direct-impact weapon, bringing the tail-as-weapon hypothesis into doubt.

Dental microwear analyses of ungulates and other large herbivores rely on correlations of diet and microwear among extant ungulates, primarily ruminants. Microwear is considered a ‘taxon-free’ method of paleodietary analysis. The properties of food are associated with causality of microwear, but the possibility that heritable properties of the consumer (tooth morphologies, masticatory dynamics, enamel mechanical properties, digestive physiologies) may introduce bias is not considered. Using an observer blind method of light microscopy, we examined the distribution of microwear features on the molars of eight species of ruminants and perissodactyls. Grazing and browsing ruminants had statistically different numbers of scratches forming discrete data clusters. Perissodactyls differ in the numbers of scratches and pits but without discrete browser and grazer clusters. Microwear features were distributed homogeneously across ruminant molars and strongly predictive of diet from the labial edge of the molar to the lingual edge. Microwear was heterogeneously distributed across perissodactyl molars with more pits on the labial edge and more scratches on the lingual edge. In perissodactyls, microwear sampled from the labial edge was strongly predictive of diet, while microwear sampled from other areas were not. Discriminant function analyses of microwear assigned individual molars to diets (browser and grazer) and clades (ruminant and perissodactyl) with similar success (70–73%) indicating that phylogeny and diet influence microwear equally. Rhino microwear was more sensitive to clade membership while other perissodactyl microwear was more sensitive to diet. Although it is not clear what heritable variables may phylogenetically bias dental microwear, extant ruminants may not be appropriate models for the microwear of other large herbivores.

Functional consequences of the variation in geometry and morphology of the articulate brachiopod hinge mechanism are poorly understood, despite the fact that hinge structures have considerable importance in brachiopod taxonomy. Jaanusson (1971) proposed that the ability to resorb shell material during growth, particularly in the hinge structures, can be used to distinguish two groups within the articulates, the deltidiodonts and the cyrtomatodonts. He considered the two groups to be morphologically distinct, “natural” phylogenetic groups, separated by a “functional discontinuity.” In order to test the morphological, functional, and phylogenetic implications of shell resorption, comparisons of the hinge-system geometry and diductor muscle moment are made here between deltidiodont and cyrtomatodont brachiopods. A truss network composed of landmarks relevant to the valve opening mechanism is constructed to characterize hinge-system geometry. The function of the hinge mechanism is analyzed in the context of valve opening, and diductor muscle force, effort lever arm, and moment are compared between deltidiodonts and cyrtomatodonts. The distribution of resorption among brachiopods is investigated with respect to a phylogenetic hypothesis proposed by Williams and Rowell (1965a).

Deltidiodont brachiopods are morphologically more variable than cyrtomatodonts, and a greater proportion of the variability is correlated with size. Deltidiodonts and cyrtomatodonts employ different strategies to open the valves; deltidiodont lever arms are relatively longer, whereas cyrtomatodont diductor muscles have relatively larger cross-sectional areas. The greatest muscle moment in deltidiodont hinge systems is realized in the maintenance of a gape angle; in the cyrtomatodont system, it is achieved at the initiation of a gape. Although they are morphologically and functionally distinct, it is doubtful that the two groups are separated by a “functional discontinuity.” Because the phylogenetic relationships among brachiopod orders are not yet resolved, the status of shell resorption as a homologue is still unclear. Resorption is manifest in at least some members of each major group of articulates (except the pentameraceans); it is likely that resorption has evolved independently several times in brachiopod evolution, in part because of the increased morphological flexibility it confers.