Drillholes made by naticid and muricid gastropods are frequently used in evolutionary and ecological studies because they provide direct, preservable evidence of predation. The muricid Ecphora is common in many Neogene Atlantic Coastal Plain assemblages in the United States, but is frequently ignored in studies of naticid predation. We used a combination of Pliocene fossil, modern beach, and experimentally derived samples to evaluate the hypothesis that Ecphora was an important source of drillholes in infaunal bivalve prey shared with naticids. We focused on the large, thick-shelled venerid, Mercenaria, which is commonly drilled by naticids today. Laboratory experiments, modern beach samples, and the published literature confirm that naticids preferentially drill near the umbo (significant clumping of holes), show a significant correlation between prey size and predator size (estimated by outer borehole diameter), and prefer Mercenaria <50 mm antero-posterior width when other prey are present. Fossil samples containing Ecphora (with or without other large muricids) show no drillhole site stereotypy (no significant clumping, greater variability in placement), no significant predator: prey size correlation, drilled prey shells larger than the largest modern naticids could produce in an experimental setting, and drillholes larger in diameter than those estimated for the largest Pliocene naticids, thus supporting our hypothesis. Substantial overlap in the placement of holes drilled by naticids and muricids, however, made identifying predators from drillhole position problematic. The lack of overlapping ranges of prey shell thickness between fossil and other samples precluded the use of drillhole morphology to establish predator identity (e.g., ratio of inner borehole diameter to outer borehole diameter, drillhole angle). Whereas the difficulty in determining predator identity from drillholes limits the types of analyses that can be reliably performed in mixed-predator assemblages, recognizing Ecphora as a prominent drilling predator creates the opportunity to investigate previously unrecognized questions.

The gomphotheres were a diverse and widespread group of proboscideans occupying Eurasia, North America, and South America throughout the Neogene. Their decline was temporally and spatially heterogeneous, and the gomphotheres ultimately became extinct during the late Pleistocene; however, the genus Cuvieronius is rarely represented in late Pleistocene assemblages in North America. Two alternative hypotheses have been invoked to explain this phenomenon: (1) competitive exclusion by sympatric mammoths and mastodons or (2) ecologic displacement due to an environmental transition from closed forests to open grasslands. To test whether competition for resources contributed to the demise of North American Cuvieronius, we present herein a large collection of stable isotope and dental microwear data from populations occupying their Pleistocene refugium in the Atlantic Coastal Plain. Results suggest that Cuvieronius consumed a wide range of resources with variable textural and photosynthetic properties and was not specialized on either grasses or browse. Further, we document evidence for the consumption of similar foods between contemporaneous gomphotheres, mammoths, and mastodons. The generalist feeding strategy of the gomphotheres likely facilitated their high Miocene abundance and diversity. However, this “jack of all trades and master of none” feeding strategy may have proved challenging following the arrival of mammoths and likely contributed to the extirpation of Cuvieronius in North America.

Barnacles living along rocky shores provide the classic example of competitive dominance in the marine ecosystem: by means of firm attachment and rapid growth, balanoid barnacles commonly undercut and overgrow the genus Chthamalus, which is thereby restricted to the upper fringe of the intertidal zone, where balanoids are physiologically incapable of living. Today, after perhaps less than 50 Myr of evolution, balanoid barnacles are in the midst of rampant adaptive radiation, being represented by about 273 species, of which about half are free-living species of intertidal or shallow subtidal habitats. Chthamaloid barnacles, in contrast, are on the decline, having originated at least 70 Myr ago but today comprising only about 53 living species, approximately 40 of which occupy the uppermost intertidal. The remainder persist as localized, relict, and often disjunct populations. Through competitive exclusion, balanoid barnacles have apparently caused the ecological restriction and decline of the chthamaloids. The balanoids have an advanced feeding mechanism, but the most important adaptive breakthrough leading to their competitive success was probably the origin of a tubiferous wall structure, which affords rapid skeletal growth for the efficient monopolization of free space and for the destruction of chthamaloids.

Nonparametric randomization tests were used to determine whether fossil marine families classified in different higher taxa are significantly different in their durations, their rates of origination and extinction, and their proportion of originations. For each of these properties, significant differences among orders and classes were found that could not be explained by differences at other taxonomic ranks, but no such differences were found among phyla. Thus, orders and classes appear to possess an evolutionary coherence that phyla lack.

Persistence in the structure of ecological communities can be predicted both by deterministic and by stochastic theory. Evaluating ecological patterns against the neutral theory of biodiversity provides an appropriate methodology for differentiating between these alternatives. We traced the history of benthic foraminiferal communities from the Huon Peninsula, Papua New Guinea. From the well-preserved uplifted reef terrace at Bonah River we reconstructed the benthic foraminiferal communities during a 2200-year period (9000–6800 yr B.P.) of reef building during the Holocene transgressive sea-level rise. We found that the similarity of foraminiferal communities was consistently above 60%, even when comparing communities on either side of a massive volcanic eruption that smothered the existing reef system with ash. Similarly, species diversity and rank dominance were unchanged through time. However, similarity dropped dramatically in the final stages of reef growth, when accommodation space was reduced as sea-level rise slowed. We compared the community inertia index (CII) computed from the observed species abundances with that predicted from neutral theory. Despite the differences in foraminiferal community composition in the younger part of the reef sequence, we found an overall greater degree of community inertia with less variance in observed communities than was predicted from neutral theory, regardless of foraminiferal community size or species migration rate. Thus, persistent species assemblages could not be ascribed to neutral predictions. Ecological incumbency of established foraminiferal species likely prevented stochastic increases in both migrant and rare taxa at the Bonah River site. Regardless of the structuring mechanisms, our reconstruction of Holocene foraminiferal assemblages provides historical context for the management and potential restoration of degraded species assemblages.

The rates and patterns of phyletic body size increase and decrease are discussed on the assumption that size change proceeds toward some limit due to selective advantage. Because the selection pressure must decrease as the average body size of an evolving population approaches the limit, some sigmoidal curve should be regarded as more appropriate for the model of size increase than an exponential curve as in the case of population growth. For phyletic size decrease, the predicted pattern is similar to a radiometric decay curve. The conventional unit of morphological rate, darwin, has been used on the assumption that the change is exponential, but actual lineages in the fossil record may represent only fractions of such sigmoidal curves. Some actual data on size increases in ceratopsian dinosaurs and Jurassic bivalves are examined, and it is concluded that sigmoidal size increase seems also empirically to be a more widespread pattern than exponential.

Relaxed clock models are fundamental in Bayesian clock dating, but a single distribution characterizing the clock variation is typically selected. Hence, I developed a new reversible-jump Markov chain Monte Carlo (rjMCMC) algorithm for drawing posterior samples between the independent lognormal (ILN) and independent gamma rates (IGR) clock models. The ability of the rjMCMC algorithm to infer the true model was verified through simulations. I then applied the algorithm to the Mesozoic bird data previously analyzed under the white noise (WN) clock model. In comparison, averaging over the ILN and IGR models provided more reliable estimates of the divergence times and evolutionary rates. The ILN model showed slightly better fit than the IGR model and much better fit than the autocorrelated lognormal (ALN) clock model. When the data were partitioned, different partitions showed heterogeneous model fit for ILN and IGR clocks. The implementation provides a general framework for selecting and averaging relaxed clock models in Bayesian dating analyses.

Investigation of the evolution of calcareous nannoplankton is hindered by the extremely small size (5-10 μm) of their fossils. We introduce new technology, a scanning electron microscope connected to an image analysis system (SEM-IAS), which allows this field to be explored as never before. This system enables a host of morphocharacters to be measured and included in studies of phylogeny and evolutionary dynamics. We have applied the SEM-IAS to study the evolution of Paleocene coccolith genera Cruciplacolithus, Chiasmolithus, and Sullivania. A variety of detailed measurements have been made on over 4000 coccoliths from Deep Sea Drilling Project Site 384 in the temperate North Atlantic and Ocean Drilling Program Site 690 in the Weddell Sea off Antarctica. Our results indicate no simple relationships between morphocharacters, the shapes of all three genera are both complex and highly variable. Although most morphocharacters possess little phylogenetic significance, the areas of different shield cycles show gradual divergence between Chiasmolithus and Sullivania through the Paleocene. Change of most other morphocharacters occurs at variable rates and reversals in trends are common. Minimal correlation exists between the trends and oscillatory shape changes observed at the two sites. We conclude that these trends and oscillations represent local, transitory ecophenotypic variation of the complex form. There is little stasis in the ten-million-year record studied.

Evolving interactions between predators and prey constitute one of the major adaptive influences on marine animals during the Paleozoic. Crinoids and fish constitute a predator–prey system that may date back to at least the Silurian, as suggested by patterns of crinoid regeneration and spinosity in concert with changes in the predatory fauna. Here we present data on the frequency of breakage and regeneration in the spines of the Middle Devonian camerate Gennaeocrinus and late Paleozoic cladids, as well as an expanded survey of the prevalence of spinosity and infestation by platyceratid gastropods on crinoid genera during the Paleozoic. Spine regeneration frequency in the measured populations is comparable to arm regeneration frequencies from Mississippian Rhodocrinites and from modern deep-water crinoid populations. The prevalence of spinosity varies by taxon, time, and anatomy among Paleozoic crinoids; notably, spinosity in camerates increased from the Silurian through the Mississippian and decreased sharply during the Pennsylvanian, whereas spines were uncommon in cladids until their Late Mississippian diversification. Among camerates, tegmen spinosity is positively correlated with the presence of infesting platyceratid gastropods. These results allow us to evaluate several hypotheses for the effects of predation on morphological differences between early, middle, and late Paleozoic crinoid faunas. Our data corroborate the hypothesis that predators targeted epibionts on camerate crinoids and anal sacs on advanced cladids and suggest that the replacement of shearing predators by crushing predators after the Hangenberg extinction affected the locations of spines in Mississippian camerates.

A hierarchical birth-death process is often used as a model for the diversification of species and higher taxa. We evaluate the validity of this null hypothesis by introducing a new measure: the distribution of species within higher taxa. The proportion of monotypic taxa and the number of species in the largest taxon generated by 1000 simulations are compared with extant mammalian data. The initial model is amended to include two sampling patterns, polyphyletic origins as well as ecological and genetic constraints on higher taxon origination rates. Simulated results are extremely variable. For genera, they are found to predict the true distribution of species quite well. For families, however, the null hypothesis is rejected in all its forms. Simulated distributions have both too many monotypic taxa and too large a dominant taxon. The latter finding stands in contrast to previous claims. Interestingly, although polyphyly and constraints slightly improve the fit of the simulations to the data, sampling does not. Finally, Smith and Patterson's dismissal of monotypic taxa is reviewed in light of this model. We argue that while some apparently monotypic taxa are sampling artifacts, this observation has no bearing on the true proportion of monotypic taxa.

Long-term persistence of patterns in geographic variation within species is an interesting and puzzling phenomenon. I present a well-defined natural experiment in the land snail Cerion agassizi from the islands of Great Bahama Bank. C. agassizi is the best-known fossil of the ca. 120,000 years BP dunes of New Providence, Cat and Eleuthera Islands; populations have survived on Cat and Eleuthera. During the Wisconsin glacial advance, all these islands joined together in an emergent bank. Presence of the same species on two islands at two times permits a test for both time signatures (does change occur in the same manner on both islands) and island signatures (do aspects of shell phenotypes remain constant on each island through time).

Factor and discriminant analyses establish morphological separations among fossil populations of the three islands. These differences occur along pathways specified by well-known covariance sets in the complex allometric ontogeny of Cerion. By these routes, small variations in the geometry of growth may be magnified to large differences in external appearance. I found a time signature, probably attributable to introgression of modern populations by Cerion glans on both Cat and Eleuthera. Despite the intermediate period of emergence and joining of all islands, I also found an island signature in the preservation through time, on both Cat and Eleuthera, of the differentia that separate fossil populations. The basic distinctions of the two islands, expressed as patterns of covariance in growth, have been stable for at least 120,000 years.

Prominent extinction “events” have been recognized from statistical analyses of marine invertebrate genera represented in Mesozoic and Cenozoic assemblages, contrasting with relatively low “background” extinction intensities measured in terms of a “percentage extinction” index. On a logarithmic scale, the slope of the relationship between time and extinction intensity for background extinctions is shown to be parallel to the slope obtained for most extinction events, characterized by intensities 100.35 above prevailing background levels. Although extinction intensities are variable, this study suggests that the magnitude of the factor(s) primarily associated with most mass extinctions in a 260-m.y. period (N = 9) need not necessarily have been very different from one event to another, an exception being the mass extinction at the end of the Cretaceous.

The mode of locomotion of any swimming animal is constrained by its size, architecture, and phylogenetic history. Considering these factors and the range of locomotory systems used by extant zooplankton, the range of possible modes of locomotion for graptoloids can be effectively limited. Three assumptions have been made: (1) graptoloids did not use a mode of locomotion unknown among modern organisms; (2) all graptoloids employed essentially the same mode of locomotion except, possibly, in their early growth stages; and (3) graptoloids did not rely entirely on passive buoyancy—no extant zooplankton groups in the size range of the graptoloids do. Structures that increase buoyancy or drag are often found in actively swimming zooplankton. They enhance feeding efficiency and reduce sinking rates during nonswimming periods.

The modes of locomotion utilized by extant zooplankton groups are ciliary propulsion, elongate body undulation, jet propulsion, rowing with skeletonized appendages, and rowing or undulation with muscular appendages. Of these, all but the last can be rejected for the graptoloids on the basis of scale or architecture. It is concluded that graptoloids probably used a rowing or undulatory motion with muscular appendages for swimming. Using a pterobranch model for the graptoloid zooids, the lophophore is considered an unlikely propulsive structure because the design requirements would conflict with those of a ciliarly suspension-feeding organ. Winglike, lateral extensions of the muscular cephalic shield, the same structure used for creeping locomotion in the benthic pterobranchs, is regarded as the most likely propulsive organ, analogous to the pteropod swimming wings.

Under stress, corals and foraminifera may eject or consume their algal symbionts (“bleach”), which can increase mortality. How bleaching relates to species viability over warming events is of great interest given current global warming. We use size-specific isotope analyses and abundance counts to examine photosymbiosis and population dynamics of planktonic foraminifera across the Paleocene–Eocene thermal maximum (PETM, ~56 Ma), the most severe Cenozoic global warming event. We find variable responses of photosymbiotic associations across localities and species. In the NE Atlantic (DSDP Site 401) PETM, photosymbiotic clades (acarininids and morozovellids) exhibit collapsed size-δ13C gradients indicative of reduced photosymbiosis, as also observed in Central Pacific (ODP Site 1209) and Southern Ocean (ODP Site 690) acarininids. In contrast, we find no significant loss of size-δ13C gradients on the New Jersey shelf (Millville) or in Central Pacific morozovellids. Unlike modern bleaching-induced mass mortality, populations of photosymbiont-bearing planktonic foraminifera increased in relative abundance during the PETM. Multigenerational adaptive responses, including flexibility in photosymbiont associations and excursion taxon evolution, may have allowed some photosymbiotic foraminifera to thrive. We conclude that deconvolving the effects of biology on isotope composition on a site-by-site basis is vital for environmental reconstructions.

We document a positive and strong correlation between speciation and extinction rates in the Paleozoic zooplankton graptoloid clade, between 481 and 419 Ma. This correlation has a magnitude of ~0.35–0.45 and manifests at a temporal resolution of <50 kyr and, for part of our data set, <25 kyr. It cannot be explained as an artifact of the method used to measure rates, sampling bias, bias resulting from construction of the time series, autocorrelation, underestimation of species durations, or undetected phyletic evolution. Correlations are approximately equal during the Ordovician and Silurian, despite the very different speciation and extinction regimes prevailing during these two periods, and correlation is strongest in the shortest-lived cohorts of species.

We infer that this correlation reflects approximately synchronous coupling of speciation and extinction in the graptoloids on timescales of a few tens of thousands of years. Almost half of graptoloid species in our data set have durations of <0.5 Myr, and previous studies have demonstrated that, during times of background extinction, short-lived species were selectively targeted by extinction. These observations may be consistent with the model of ephemeral speciation, whereby new species are inferred to form constantly and at high rate, but most of them disappear rapidly through extinction or reabsorption into the ancestral lineage. Diversity dependence with a lag of ~1 Myr, also documented elsewhere, may reflect a subsequent and relatively slow, competitive dynamic that governed those species that dispersed beyond their originating water mass and escaped the ephemeral species filter.

Pollen malformations have been proposed as a paleoenvironmental stress proxy. However, the frequency and variability of pollen malformations under near-optimal conditions and environmental stress, as well as their developmental origins, remain unclear. To bridge these gaps, we compared pollen malformation frequencies and assemblages of 14 extant conifer genera of Pinaceae and Podocarpaceae producing saccate (winged) grains grown under near-optimal conditions. These baseline pollen yields were compared with those produced by Pinus mugo ‘Columnaris’ cultured under an abiotic stress—three experimentally heightened ultraviolet-B radiation (UV-B) regimes proposed for the end-Permian crisis. We additionally reviewed previous cytological literature of abnormal microsporogenesis in conifers. Under near-optimal conditions, malformations comprise <3% of pollen yields in 12 out of 13 bisaccate genera and >10% of yields in the naturally trisaccate Dacrycarpus dacrydioides. We detected no phylogenetic pattern in malformation assemblages of the baseline comparisons. UV-B–irradiated P. mugo produced significantly higher malformation frequencies and different assemblage compositions when compared with baseline bisaccate lineages. We propose that pollen malformations originate during the meiotic and tetrad stages of microsporogenesis and present a framework for the ontogeny of different malformation types seen in the fossil record. Malformations comprising >3% of bisaccate pollen yields can be used as a paleoenvironmental stress proxy, but rare, naturally trisaccate lineages are not suitable for such assessments. Furthermore, heightened UV-B not only increases pollen malformation production, but also alters the types of abnormalities trees produce. Different environmental stresses may therefore leave behind distinct fingerprints in the fossil record.

The geometry of the simpler types of graptolite stipes can be quantified by assuming them to consist of stacked cylinders representing the thecal tubes. It can be shown that specification of any three characters, for example, thecal inclination, thecal spacing, and stipe width, effectively defines the others (thecal overlap, thecal length and width, thecal “density”); hence the parameters commonly used in the definition of species are not independent. Variation within and between species can be represented by three-character plots. The effects of altering the apertural angle from 90° on such characters as thecal spacing can also be represented geometrically. If thecae are considered as cones rather than cylinders, it follows that as stipe width increases, thecal curvature will describe a sine curve if the thecae are to remain in contact; this observation is matched on real graptolites. Thecal curvature beyond 90° to the dorsal wall is generally impossible without thecae detaching from their neighbors to produce thecal isolation. Growth of thecae along a stipe can be represented graphically. Computer fits to growth curves for five dichograptids show that they can be closely described by an exponential decrease in growth increments along the stipe, and that this model is more accurate than two possible alternatives.

Most evolutionary innovations—power-enhancing phenotypes previously absent in a lineage—have arisen multiple times within major clades. This repetition permits a comparative approach to ask how, where, when, in which clades, and under which circumstances adaptive innovations are acquired and secondarily lost. I use new and literature-based data on the phylogeny, functional morphology, and fossil record of gastropods to explore the acquisition and loss of the siphonal canal and its variations in gastropods. The siphonal indentation, canal, notch, or tube at the front end of the shell is associated in living gastropods with organs that detect chemical signals directionally and at a distance in an anteriorly restricted inhalant stream of water.

Conservative estimates indicate that the siphonate condition arose 23 times and was secondarily lost 17 times. Four siphonate clades have undergone prodigious diversification. All siphonate gastropods have a shell whose axis of coiling lies at a low angle above the plane of the aperture (retroaxial condition). In early gastropods, the siphonal canal was short and more or less confined to the apertural plane. Later (mainly Cretaceous and Cenozoic) variations include a dorsally deflected canal, a long canal, and a closed canal. The closed siphonal canal, in which the edges join to form a tube, arose 15 times, all in the adult stages of caenogastropods with determinate growth.

Gastropods in which the siphonate condition arose were mobile, bottom-dwelling, microphagous animals. Active predaceous habits became associated with the siphonate condition in the Mesozoic and Cenozoic Purpurinidae-Latrogastropoda clade. Loss of the siphonate condition is associated with nonmarine habits, miniaturization, and especially with a sedentary or slow-moving mode of life.

The siphonate condition arose seven times each during the early to middle Paleozoic, the late Paleozoic, and the early to middle Mesozoic, and only once each during the Late Cretaceous and Cenozoic. Well-adapted incumbents prevented most post-Jurassic clades from evolving a siphonal indentation and its associated organs. Dorsally deflected, long, and closed canals are known only from Cretaceous and Cenozoic marine gastropods, and represent improvements in sensation and passive armor.

In a discussion of contrasting ecologies of clades that gained and lost the siphonate condition, I argue that macroevolutionary trends in the comings and goings of innovations and clades must incorporate ecological and functional data. Acquisitions of energy-intensive, complex innovations that yield greater power have a greater effect on ecosystems, communities, and their resident clades than do reversals, which generally reflect energy savings.