The role of antagonistic organismal interactions in the production of long-term macroevolutionary trends has been debated for decades. Some evidence seems to suggest that temporal trends in predation frequency share a common causative mechanism with genus-level diversity, whereas studies on the role of parasites in “shaping” the evolutionary process are rare indeed. Digenean trematodes (Phylum Platyhelminthes) infest molluscs in at least one stage of their complex life cycle. Trematodes leave characteristic oval-shaped pits with raised rims on the interior of their bivalve hosts, and these pits are preserved in the fossil record. Here we survey 11,785 valves from the Pleistocene–Holocene deposits of the Po Plain and from nearby modern coastal environments on the northeast Adriatic coast of Italy. Of these, 205 valves exhibited trematode-induced pits. Trematodes were selective parasites in terms of host taxonomy and host body size. Infestation was restricted to lower shoreface/transition-to-platform paleoenvironments. During the Holocene, individuals from the transgressive systems tract were significantly more likely to be infested than those from highstand systems tracts. Temporal trends in infestation frequency cannot be explained as an ecological/evolutionary phenomenon (e.g., the hypothesis of escalation); instead the trend seems controlled by environmental variation induced by glacio-eustatic sea-level changes and inadequate sampling. Because this interaction appears to be ephemeral, both temporally and spatially, it is not likely that any selective pressure would be continuous over geologic time in this region. Furthermore, these results support the hypothesis that antagonistic interactions are lower in the northern Adriatic Sea in comparison to other midlatitude shallow marine settings.

The two major approaches to studying macroevolution in deep time are the fossil record and reconstructed relationships among extant taxa from molecular data. Results based on one approach sometimes conflict with those based on the other, with inconsistencies often attributed to inherent flaws of one (or the other) data source. Any contradiction between the molecular and fossil records represents a failure of our ability to understand the imperfections of our data, as both are limited reflections of the same evolutionary history. We therefore need to develop conceptual and mathematical models that jointly explain our observations in both records. Fortunately, the different limitations of each record provide an opportunity to test or calibrate the other, and new methodological developments leverage both records simultaneously. However, we must reckon with the distinct relationships between sampling and time in the fossil record and molecular phylogenies. These differences impact our recognition of baselines and the analytical incorporation of age estimate uncertainty.

Sharks have a long and rich fossil record that consists predominantly of isolated teeth due to the poorly mineralized cartilaginous skeleton. Tiger sharks (Galeocerdo), which represent apex predators in modern oceans, have a known fossil record extending back into the early Eocene (ca. 56 Ma) and comprise 22 recognized extinct and one extant species to date. However, many of the fossil species remain dubious, resulting in a still unresolved evolutionary history of the tiger shark genus. Here, we present a revision of the fossil record of Galeocerdo by examining the morphological diversity and disparity of teeth in deep time. We use landmark-based geometric morphometrics to quantify tooth shapes and qualitative morphological characters for species discrimination. Employing this combined approach on fossil and extant tiger shark teeth, our results only support six species to represent valid taxa. Furthermore, the disparity analysis revealed that diversity and disparity are not implicitly correlated and that Galeocerdo retained a relatively high dental disparity since the Miocene despite its decrease from four to one species. With this study, we demonstrate that the combined approach of quantitative geometric morphometric techniques and qualitative morphological comparisons on isolated shark teeth provides a useful tool to distinguish between species with highly similar tooth morphologies.

The schizochroal eye of trilobites bears lenses of high optical quality. Cooperation between adjacent lenses, which must have had retinas rather than rhabdomeres, might have allowed the trilobite stereoscopic vision throughout its visual field. We propose a simple neurophysiological model for the schizochroal eye in which neural connections between lenses on vertical “visual strips” could have provided the integration necessary for stereoscopic vision. Simple neural signals could have coded information on the position, size, speed and nature of an object in the field of view. The model of stereoscopic vision is consistent with data on the shape of the eye, the arrangement of lenses on its surface, and with current views on the origin and early evolution of the schizochroal eye. It implies that the lens arrangement, neural network and shape of the eye surface were a coadapted morphological complex, and it is consistent with accepted ideas of the Phacopina as living in benthic habitats in the photic zone.

Widespread concern about environmental pollution is putting a new question to the fossil record: How has the biosphere reacted to chemical changes in the past? Monera and Protoctista might be expected to provide valuable clues in this quest since their biomineral remains are generally formed in conditions closely related to the environment. But why do unicells biomineralize at all? It was with such questions in mind that an international symposium of the Systematics Association on “Biomineralization in Lower Plants and Animals” was held at Birmingham on April 15–19, 1985. Monerans, protoctistans, lichens, calcareous algae, and bryozoans were discussed in 36 papers, of which 23 are to be published in a volume by Oxford University Press. This volume, edited by Leadbeater and Riding (1986), will form a natural sequel to the papers in Miller et al. (1984) on mineral phases in biology and in Westbroek and de Jong (1983) on biomineralization and biological metal accumulation.

Gould (1980) claims that a “new and general theory of evolution” is emerging. He examines the modern synthesis and asserts that it is being rejected by present day evolutionary biologists. The purpose of this note is to show that Gould's view of the modern synthesis is distorted. Any rejection of the modern synthesis must rest on a more balanced characterization than that presented by Gould.

Muscular articulations on the brachials and pinnules of Paleozoic crinoids have traditionally been inferred by the presence of a transverse ridge. We here provide evidence that the microstructure of the stereom, fine meshed where muscles once inserted and coarse meshed where ligaments were emplaced, provides a reliable microstructural criterion for muscular articulation on well-preserved arm ossicles. Partially pyritized ossicles of Aesiocrinus magnificus Miller & Gurley from the Lane Shale of Pennsylvanian age at Kansas City, Missouri exhibit well-preserved stereom in three dimensions under the scanning electron microscope. These brachial and pinnular plates show clearly the distinctiveness of stereom at sites of muscule and ligament attachment. This criterion should be useful in tracing the evolutionary development of flexible arm articulations in several lineages of Paleozoic crinoids.

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.

The modern gorgonocephalid ophiuroid Astrophyton, a basketstar, has feeding habits which may be analogous to those of some Paleozoic crinoids that lack pinnulate arms (Meyer and Lane 1975). Field study has demonstrated the very quick response of Astrophyton when plankton impinge upon its arms. Study of the skeletal microstructure by scanning electron microscopy has shown that the quick response is facilitated by the development of relatively large ventral muscle fossae and articular surfaces whose shape follows a catenary curve. Prey are impaled by spines which articulate with the ventral surface of vertebral ossicles. Stereomic microstructure of vertebral ossicles is galleried where ligaments are present and labyrinthic where muscular tissue is present. The same microstructures are found in muscular articulations of modern crinoids. Future work on the arm structure of Paleozoic taxocrinids and cyathocrinids may demonstrate structural analogies and suggest similarities in food capture between such crinoids and modern baskets tars.

Paleobiological diversity is often expressed as α (within-sample), β (among-sample), and γ (total) diversities. However, when studying the effects of extinction on diversity patterns, only variations in α and γ diversities are typically addressed. A null model that examines changes in β diversity as a function of percent extinction is presented here.

The model examines diversity in the context of a hierarchical sampling strategy that allows for the additive partitioning of γ diversity into mean α and β diversities at varying scales. Here, the sampling hierarchy has four levels: samples, beds, facies, and region; thus, there are four levels of α diversity (α1, α2, α3, α4) and three levels of β diversity (β1, β2, and β3). Taxa are randomly assigned to samples within the hierarchy according to probability of occurrence, and initial mean α and β values are calculated. A regional extinction is imposed, and the hierarchy is resampled from the remaining extant taxa. Post-extinction mean α and β values are then calculated.

Both non-selective and selective extinctions with respect to taxon abundance yield decreases in α, β, and γ diversities. Non-selective extinction with respect to taxon abundance shows little effect on diversity partitioning except at the highest extinction magnitudes (above 75% extinction), where the contribution of α1 to total γ increases at the expense of β3, with β1 and β2 varying little with increasing extinction magnitude. The pre-extinction contribution of α1 to total diversity increases with increased probabilities of taxon occurrence and the number of shared taxa between facies. Both β1 and β2 contribute equally to total diversity at low occurrence probabilities, but β2 is negligible at high probabilities, because individual samples preserve all the taxonomic variation present within a facies. Selective extinction with respect to rare taxa indicates a constant increase in α1 and constant decrease in β3 with increasing extinction magnitudes, whereas selective extinction with respect to abundant taxa yields the opposite pattern of an initial decrease in α1 and increase in β3. Both β1 and β2 remain constant with increasing extinction for both cases of selectivity. By comparing diversity partitioning before and after an extinction event, it may be possible to determine whether the extinction was selective with respect to taxon abundances, and if so, whether that selectivity was against rare or abundant taxa.

Field data were collected across a Late Ordovician regional extinction in the Nashville Dome of Tennessee, with sampling hierarchy similar to that of the model. These data agree with the abundant-selective model, showing declines in α, β, and γ diversities, and a decrease in α1 and increase in β3, which suggests this extinction may have targeted abundant taxa.

The final 10 Myr of the Paleozoic saw two of the biggest biological crises in Earth history: the middlePermian extinction (often termed the Guadalupian–Lopingian extinction [GLE]) that was followed 7–8 Myr later by Earth's most catastrophic loss of diversity, the Permian–Triassic mass extinction (PTME). These crises are not only manifest as sharp decreases in biodiversity and—particularly for the PTME—total ecosystem collapse, but they also drove major changes in biological morphological characteristics such as the Lilliput effect. The evolution of test size among different clades of foraminifera during these two extinction events has been less studied. We analyzed a global database of foraminiferal test size (volume) including 20,226 specimens in 464 genera, 98 families, and 9 suborders from 632 publications. Our analyses reveal significant reductions in foraminiferal mean test size across the Guadalupian/Lopingian boundary (GLB) and the Permian/Triassic boundary (PTB), from 8.89 to 7.60 log10 μm3 (lg μm3) and from 7.25 to 5.82 lg μm3, respectively. The decline in test size across the GLB is a function of preferential extinction of genera exhibiting gigantism such as fusulinoidean fusulinids. Other clades show little change in size across the GLB. In contrast, all Lopingian suborders in our analysis (Fusulinina, Lagenina, Miliolina, and Textulariina) experienced a significant decrease in test size across the PTB, mainly due to size-biased extinction and within-lineage change. The PTME was clearly a major catastrophe that affected many groups simultaneously, and the GLE was more selective, perhaps hinting at a subtler, less extreme driver than the later PTME.

Miniaturization has been defined as the evolution of extremely small adult size in a lineage. It does not simply imply the decrease of the body size but also involves structural modifications to maintain functional efficiency at a strongly reduced size. Miniaturization has been proposed as a key factor in the origin of several major tetrapod clades. Current hypotheses propose that the living amphibians (lissamphibians) originated within a clade of Paleozoic dwarfed dissorophoid temnospondyls. Morphological traits shared by these small dissorophoids have been interpreted as resulting from constraints imposed by the extreme size reduction, but these statements were based only on qualitative observations. Herein, we assess quantitatively morphological changes in the skull previously associated with miniaturization in the lissamphibian stem lineage by comparing evolutionary and ontogenetic allometries in dissorophoids. Our results show that these features are not comparable to the morphological consequences of extreme size reduction as documented in extant miniature amphibians, but instead they resemble immature conditions of larger temnospondyls. We conclude that the truncation of the ancestral ontogeny, and not constraints related to miniaturization, might have been the factor that played a major role in the morphological evolution of small dissorophoids. Based on our results, we discuss the putative role of miniaturization in the origin of lissamphibians within Dissorophoidea.

The phenomenon of selective biotic extinction is commonly explained as the result of the extinction of specialized organisms and the survival of generalized organisms during a time of environmental stress. Using 48 orders of aquatic free-living arthropods, we test the hypothesis that generalized organisms tend to have longer geologic ranges than specialized organisms. The degree of specialization, as expressed through morphological complexity, is measured by the diversity of limb pairs on the arthropod body. The resulting “tagmosis value” reflects the functional differentiation of limb pairs to perform specific tasks and thus may estimate ecological specialization. No statistically significant correlation exists between an order's typical degree of morphological specialization and its geologic range. Degree of specialization is not an important determinant of evolutionary persistence at this taxonomic level. Evolutionary longevity may depend on the adaptive type (e.g., trophic role, relationship to substrate, etc.) represented by the taxon and on the assortment of these adaptive types.

Patterns preserved in the fossil record are of the highest importance in addressing questions about long-term evolutionary processes, yet both the description of pattern and its translation into process can be difficult. With respect to gradual phyletic change, we know that randomly generated sequences may exhibit characteristics of a “trend” apparent patterns, therefore, must be interpreted with caution. Furthermore, even when the claim of a gradual trend can be statistically justified, interpretation of the underlying mechanisms may be challenging. Given that we can observe populations changing rapidly over tens or hundreds of years, it is now more difficult to explain instances of geologically gradual (as opposed to punctuated) change.

Here we describe morphologic change in two bivalve lineages from the late Miocene Lake Pannon. We evaluate change according to the model-based methods of Hunt. Both lineages exhibit size increases and shape changes over an interval of nearly 4 million years. Size and two shape variables in the conjungens lineage are best fit by a model of directional evolution; remaining shape variables mostly conform to unbiased random walks. Body-size evolution in the diprosopum lineage is also significantly directional but all shape variables are best fit by the unbiased random walk model; the small number of sampling intervals available for this lineage (n = 6) makes determination of the actual pattern more difficult. Model-fitting results indicate that the parallel trajectories of increasing log shell height over time in the two lineages can be accounted for by an underlying trend shared by both lineages, suggesting that the size increases may be a shared response to the same cause. The pace of phenotypic change, measured as Lynch's Δ, is slower than the neutral expectation for all size and shape traits.

Our examples illustrate well the paradox of gradualism; the sequences exhibit significant directional morphological evolution, but rates of change as measured over the long-term are apparently too slow for directional selection or even drift to be the cause. Viewing long-term phenotypic evolution in terms of populations tracking peaks on adaptive landscapes is useful in this context. Such a view allows for intervals of directional selection (during times of peak movement–resulting in the overall trends we can detect) interspersed with intervals of stasis (during times of peak stability–resulting in overall changes that appear to proceed more slowly than the neutral expectation). The paradox of gradualism thus reduces to (1) peak movements and their drivers, which are not restricted in rate as are population-genetic drivers, and (2) the maintenance of stasis, on which no consensus exists.

We can identify no environmental parameter in the central European Neogene that exhibits consistent change across the interval of gradual morphologic change. It may be that in Lake Pannon the long-term persistence of generally ameliorating conditions (plentiful resources and habitat space, few predators or competitors) resulted in geologically slow but consistent peak shifts, which in turn facilitated size increase and shape change in these lineages.

Steneofiber, an early Miocene European beaver, showed evidence of a K-strategy model of reproduction and important morphological modifications for better adaptation to a semiaquatic way of life. At the same time, a North American branch of Castoridae developed not only a parental care system but also a series of cranial and postcranial osteological adaptations to a burrowing way of life. As the North American and the European Castoridae evolved independently from each other for at least fifteen m.y., these behavioral trends seem to have been inherent in the family for a long time and have apparently promoted, up to the present time, the spread of semiaquatic forms on the two continents; on the contrary, the burrowing branch of the Castoridae rapidly vanished.

The late Pleistocene megafaunal extinctions may have been the first extinctions directly related to human activity, but in North America the close temporal proximity of human arrival and the Younger Dryas climate event has hindered efforts to identify the ultimate extinction cause. Previous work evaluating the roles of climate change and human activity in the North American megafaunal extinction has been stymied by a reliance on geographic binning, yielding contradictory results among researchers. We used a fine-scale geospatial approach in combination with 95 megafaunal last-appearance and 75 human first-appearance radiocarbon dates to evaluate the North American megafaunal extinction. We used kriging to create interpolated first- and last-appearance surfaces from calibrated radiocarbon dates in combination with their geographic autocorrelation. We found substantial evidence for overlap between megafaunal and human populations in many but not all areas, in some cases exceeding 3000 years of predicted overlap. We also found that overlap was highly regional: megafauna had last appearances in Alaska before humans first appeared, but did not have last appearances in the Great Lakes region until several thousand years after the first recorded human appearances. Overlap in the Great Lakes region exceeds uncertainty in radiocarbon measurements or methodological uncertainty and would be even greater with sampling-derived confidence intervals. The kriged maps of last megafaunal occurrence are consistent with climate as a primary driver in some areas, but we cannot eliminate human influence from all regions. The late Pleistocene megafaunal extinction was highly variable in timing and duration of human overlap across the continent, and future analyses should take these regional trends into account.

The occupation of new environments by evolutionary lineages is frequently associated with morphological changes. This covariation of ecotype and phenotype is expected due to the process of natural selection, whereby environmental pressures lead to the proliferation of morphological variants that are a better fit for the prevailing abiotic conditions. One primary mechanism by which phenotypic variants are known to arise is through changes in the timing or duration of organismal development resulting in alterations to adult morphology, a process known as heterochrony. While numerous studies have demonstrated heterochronic trends in association with environmental gradients, few have done so within a phylogenetic context. Understanding species interrelationships is necessary to determine whether morphological change is due to heterochronic processes; however, research is hampered by the lack of a quantitative metric with which to assess the degree of heterochronic traits expressed within and among species. Here I present a new metric for quantifying heterochronic change, expressed as a heterochronic weighting, and apply it to xiphosuran chelicerates within a phylogenetic context to reveal concerted independent heterochronic trends. These trends correlate with shifts in environmental occupation from marine to nonmarine habitats, resulting in a macroevolutionary ratchet. Critically, the distribution of heterochronic weightings among species shows evidence of being influenced by both historical, phylogenetic processes and external ecological pressures. Heterochronic weighting proves to be an effective method to quantify heterochronic trends within a phylogenetic framework and is readily applicable to any group of organisms that have well-defined morphological characteristics, ontogenetic information, and resolved internal relationships.

A number of facts indicate that the diversity of chthamaloid barnacles is much lower now than it has been in the past. Among these are the relict, often disjunct geographic distributions of distinctive chthamaloid higher taxa and their representation by only a few living species. The living chthamaloid barnacles also have a disjunct bathymetric distribution: as we will document here in detail, they are concentrated in the intertidal zone and in deep water. In his rejoinder to our paper, Paine makes no cogent argument against our interpretations of this powerful evidence.

Our case that the balanoids have competitively displaced the more primitive chthamaloids is based on several facts: (1) the Chthamaloidea have dwindled during the Cenozoic Era, so that most have relict or refugial distributions, whereas the Balanoidea have undergone a remarkable adaptive radiation; (2) balanoid species are known to defeat chthamaloid species in competition for space; (3) the competitive success of the balanoids can be attributed to an inherent biological feature—a tubiferous skeleton, which fosters rapid growth; (4) the peak diversity of living balanoids coincides precisely with the bathymetric gap in the chthamaloids’ distribution.

We argue that the radiation of balanoids during the past 40 Myr or so has caused a decline in chthamaloid diversity by reducing and destabilizing low intertidal and shallow subtidal populations, thus elevating average rate of extinction and depressing average rate of speciation. This model allows for exceptions to the rule of balanoid competitive dominance, but contrary to Paine's claim, the exceptions that exist are either trivial or support our view.

Paine offers no alternative to our competition hypothesis. He suggests that an increase in predation has caused a decline in chthamaloid body size, but neither a Cenozoic trend in body size nor one in predation pressure has empirical support; furthermore, the body-size hypothesis has no direct bearing on the case we have presented.

The hypothesis of limiting similarity, which postulates that morphologically and/or ecologically similar species will differ enough in shape, size, or other variables to minimize competition, has been controversial among ecologists and paleoecologists. Many studies have reported the occurrence of limiting similarity in modern environments or in time-averaged fossil deposits; however, empirical high-resolution time series demonstrating limiting similarity over longer time scales are lacking. We have integrated radiocarbon-calibrated amino acid dating techniques, stable isotope estimates, and morphometric data to test the hypothesis of limiting similarity in late Quaternary land snails from the Canary Islands over a period of 42,500 years. We tested for both ecological character displacement (two closely related species will differ in size in order to minimize competition in sympatry and these differences will be minimized in allopatry) and communitywide character displacement (overdispersion of body size among competitors in a guild). Multiple proxies of body size consistently show that two endemic congeneric pulmonate gastropod species (Theba geminata and T. arinagae) maintained a difference in size from ~42,500 B.P. through the last occurrence of T. arinagae 14,900 B.P., with a concomitant trend of a decreasing body size. Theba geminata body size did not converge on that of T. arinagae and variation in T. geminata body size did not increase significantly following the extinction of T. arinagae; therefore, ecological character displacement and release did not occur. Community-wide character displacement was found in only one time bin over the last 42,500 years. These results suggest that limiting similarity is a transient ecological phenomenon rather than a long-term evolutionary process. This study not only demonstrates the problems inherent in biological “snapshot” studies and geological studies of time-averaged deposits to test limiting similarity adequately, but it also presents a more adequate research protocol to test the importance of interspecific competition in the history of life.