Diet and body mass are highly important factors in mammalian ecology, and they have also proven to be powerful paleoecological indicators. Our previous research has proposed a new classification scheme for mammals with more dietary divisions that emphasizes the primary resource in a given diet. We analyzed a database summarizing the dietary preferences of 139 species of marsupial and placental terrestrial mammals (including 14 orders) and their average body masses in order to explore whether this new classification better highlights ecomorphological differences between species. Additionally, the dietary diversity of every species in the data set was quantified by applying the inverse Simpson index to stomach content percentages. We observed a decrease in maximum dietary diversity with increasing body mass. Having lower requirements for energy and nutrients per unit of body weight or ecological advantages such as larger home ranges allows larger mammals to feed on less nutritive feeding resources (i.e., structural plant material). Our results also suggest that body-size ranges are different across dietary specializations. Smaller mammals (<1 kg) are mainly insectivores, granivores, or mixed feeders, while bigger animals (>30 kg) are usually either carnivores or herbivores that feed specifically on grasses and leaves. The medium-size range (1–30 kg) is mostly composed of frugivorous species that inhabit tropical and subtropical rain forests. Thus, the near absence of medium-sized mammals in open environments such as savannas can be linked to the decreasing density of fruit trees needed to support a pure frugivorous diet year-round. In other words, seasonality of precipitation prevents species from specializing on a totally frugivorous diet. Our results suggest that this new classification scheme correlates well with body mass, one of the most studied morphological variables in paleoecology and ecomorphology. Therefore, the classification should serve as a useful basis for future paleoclimatological studies.

Ecological niche modeling (ENM) is a quantitative approach to predict species’ abiotic requirements. It is a correlative technique, requiring geographically explicit information on species occurrences and the suites of environmental conditions experienced at each occurrence point. The output of these models is a set of environmental suitability rules that can be projected geographically and through time to test biogeographic, ecologic, and evolutionary hypotheses. Although developed by biologists and used extensively in the modern, ENM is in its early stages of application to the deep-time fossil record (hence PaleoENM). In part its limited use in the fossil record thus far reflects the methodological challenge of constructing paleoenvironmental layers needed for PaleoENM analysis, whereas in the modern these layers are available from large public databases (e.g., WorldClim). This paper provides a contextual and methodological framework for appropriately applying PaleoENM, including best practices for developing species occurrence and paleoenvironmental data sets for PaleoENM analyses.

Fossil vertebrate distributions are typically based on body fossils, which are often poorly sampled at the margins of their true temporal and spatial ranges. Because vertebrate ichnofossils can be preserved in great abundance and in different environments than vertebrate body fossils, inclusion of ichnofossil data may improve sampled ranges. However, if ichnofossils are to serve as an independent source of distributional data, then their attribution to a body fossil group (i.e., trackmaker identification) cannot rely on temporal and spatial coincidence. Ichnofossils identified by synapomorphies can act as an independent source of distributional data that can modify spatial, temporal, and character distributions, which in turn may influence hypotheses of locomotor evolution.

In this paper I evaluate the spatial, temporal, and character distributions of early sauropod dinosaurs by using a combined ichnofossil and body fossil data set. Sauropod ichnofossils supplement the spatiotemporal distributions of early sauropods and provide important information on early sauropod foot posture that is rarely preserved or can only be inferred from body fossils. The presence of derived features in early-appearing ichnofossils challenges previous hypotheses of character transformation, implying either parallelism, reversal, or ghost lineages.

Stratocladistics can be used to resolve conflicting character and temporal distributions from body fossils and ichnofossils. Stratocladistic analysis of a combined ichnofossil and body fossil data set suggests a richer, more widely distributed diversity of early sauropods than currently recognized in body fossils and suggests that several locomotor characters originated much earlier than implied by body fossils.

Models of “ideal” orthoconic shells having simple concave septa with minimal weight and maximal strength and analysis of 72 species of fossil orthocones and cyrtocones yield important insights into the physical principles underlying cephalopod shell design. The ideal septum is a spherical cap weighing only 77% of a hemispherical septum of equal strength. The septa of most longicones approximate this ideal shape while those of brevicones are less curved, probably owing to buoyancy problems. Increase in septal strength leads to weight increase unless the shell becomes more logiconic or septal spacing increases or both. However, increased spacing requires more cameral liquid for septum formation, thus reducing buoyancy. In ideal longicones, septal spacing resembles the cone radius for thick, strong septa but declines to half of the cone radius for thin, weak septa. In ideal intermediates and brevicones, spacings are respectively reduced by factors of about 2 and 4, with similar additional dependence on septal thickness. Most real septa resemble these ideal models.

The relative length of the body chamber to the phragmocone varies greatly between about 0.2 and 1.5, depending mainly on the wall thickness and to a lesser degree on the septal thickness, apical angle and body density. Removal of cameral liquid in the adult must be compensated for by additional growth to retain neutral buoyancy. The conditions for neutral equilibrium calculated for longicones with different “counterweights” indicate that: (1) cameral liquid only is least feasible; (2) half-and-half calcium carbonate and liquid results in one-third length and one-quarter volume reduction of the body chamber; (3) with calcium carbonate only, body chamber reduction is minimal. Real ‘counterweights’ appear to be intermediate between (2) and (3), providing the animal with horizontal stability, which is missing in (3). Most uncalcified siphuncles reduce the body chamber only slightly although they improve horizontal stability. If the wall attains full thickness only at the apical end of body chamber, the liquid-only ‘counterweight’ becomes feasible.

The view that interspecific competition with balanoid barnacles has caused the evolutionary decline of chthamaloids (Stanley and Newman 1980), is challenged on 3 grounds. First, as a phyletic stock, chthamaloid size, as indicated by the mean adult basal diameter of extant taxa, has been decreasing, probably since the end of the Cretaceous. Small organisms will lose in interspecific competition to larger ones. However small body size reduces the susceptibility to predation and various forms of disturbance, and therefore allows chthamaloids to inhabit intertidal areas denied to balanoids. Second, although many species of Chthamalus occupy a high intertidal “refuge,” there is little evidence that competition is the primary ecological force maintaining them there. Predation is the most significant influence in tropical areas. In many temperate zone communities characterized by intense invertebrate predation or physical disturbance, chthamaloids achieve their maximum density lower in the intertidal than do balanoids or coronuloids. Last, the fossil evidence supporting a decline in chthamaloids is missing. The difficulty of seeking a single mechanism to explain global, multispecific phenomena is discussed.

The concept of coordinated stasis, manifest as a pattern of long intervals of concurrent taxonomic and ecologic persistence separated by comparatively abrupt periods of biotic change, has been challenged in recent studies that claim a lack of prolonged persistence of taxa and associations. A key problem has been the difficulty of distinguishing faunal change owing to localized, short-term environmental fluctuation or patchiness from that indicating regionally pervasive, long-term evolutionary or ecological change. Here, we use an extensive database from the Middle Devonian Hamilton Group of the Appalachian Basin to test for taxonomic and ecologic persistence within this ecological-evolutionary subunit, a succession of purported relative stability. Replicate samples collected from many localities and stratigraphic horizons over a wide geographic area allow us to address the effects of small-scale environmental variation and localized faunal patchiness while exploring basin-scale variation in faunal composition within and between the formations of the Hamilton Group.

Observed stratigraphic distributions of fossils are consistent with a scenario in which all taxa are present from bottom to top of the Hamilton Group, and absences result only from sampling failure. Although small-scale variation in faunal composition indeed does occur, there is no more variation among formations than occurs within them. Assemblages from different formations, whether they are defined by taxonomic or ecologic composition, are statistically indistinguishable according to several independent metrics, including ANOSIM and a maximum likelihood estimation that evaluates stratigraphic turnover using Bayesian “Information Criterion.” Simulated data sets indicate that test results are most consistent with species-level extinction of 2.6% per Myr within the Hamilton Group, far lower than the Givetian rate of 11.5% per Myr generic extinction derived from a global database. Such faunal persistence over the ~5.5 Myr encompassed by this unit is consistent with the pattern of coordinated stasis. Earlier studies showing greater amounts of temporal turnover in Hamilton Group faunas are likely influenced by their smaller geographic scale of analysis, suggesting that regional studies done elsewhere may yield similar results.

Although only a few studies have explicitly evaluated live-dead agreement of species and community responses to environmental and spatial gradients, paleoecological analyses implicitly assume that death assemblages capture these gradients accurately. We use nine data sets from modern, relatively undisturbed coastal study areas to evaluate how the response of living molluscan assemblages to environmental gradients (water depth and seafloor type; “environmental component” of a gradient) and geographic separation (“spatial component”) is captured by their death assemblages. We find that:

1. Living assemblages vary in composition either in response to environmental gradients alone (consistent with a species-sorting model) or in response to a combination of environmental and spatial gradients (mass-effect model). None of the living assemblages support the neutral model (or the patch-dynamic model), in which variation in species abundance is related to the spatial configuration of stations alone. These findings also support assumptions that mollusk species consistently differ in responses to environmental gradients, and suggest that in the absence of postmortem bias, environmental gradients might be accurately captured by variation in species composition among death assemblages. Death assemblages do in fact respond uniquely to environmental gradients, and show a stronger response when abundances are square-root transformed to downplay the impact of numerically abundant species and increase the effect of rare species.

2. Species' niche positions (position of maximum abundance) along bathymetric and sedimentary gradients in death assemblages show significantly positive rank correlations to species positions in living assemblages in seven of nine data sets (both square-root-transformed and presence-absence data).

3. The proportion of compositional variation explained by environmental gradients in death assemblages is similar to that of counterpart living assemblages. Death assemblages thus show the same ability to capture environmental gradients as do living assemblages. In some instances compositional dissimilarities in death assemblages show higher rank correlation with spatial distances than with environmental gradients, but spatial structure in community composition is mainly driven by spatially structured environmental gradients.

4. Death assemblages correctly identify the dominance of niche metacommunity models in mollusk communities, as revealed by counterpart living assemblages. This analysis of the environmental resolution of death assemblages thus supports fine-scale niche and paleoenvironmental analyses using molluscan fossil records. In spite of taphonomic processes and time-averaging effects that modify community composition, death assemblages largely capture the response of living communities to environmental gradients, partly because of redundancy in community structure that is inherently associated with multispecies assemblages. The molluscan data sets show some degree of redundancy as evidenced by the presence of at least two mutually exclusive subsets of species that replicate the community structure, and simple simulations show that between-sample relationships can be preserved and remain significant even when a large proportion of species is randomly removed from data sets.

Questions related to dinosaur behavior can be difficult to answer conclusively by using morphological studies alone. As a complement to these approaches, carbon and oxygen isotope ratios of tooth enamel can provide insight into habitat and dietary preferences of herbivorous dinosaurs. This approach is based on the isotopic variability in plant material and in surface waters of the past, which is in turn reflected by carbon and oxygen isotope ratios of animals that ingested the organic matter or drank the water. Thus, it has the potential to identify and characterize dietary and habitat preferences for coexisting taxa.

In this study, stable isotope ratios from coexisting hadrosaurian and ceratopsian dinosaurs of the Hell Creek Formation of North Dakota are compared for four different stratigraphic levels. Isotopic offsets between tooth enamel and tooth dentine, as well as taxonomic differences in means and in patterns of isotopic data among taxa, indicate that primary paleoecological information is preserved. The existence of taxonomic offsets also provides the first direct evidence for dietary niche partitioning among these herbivorous dinosaur taxa. Of particular interest is the observation that the nature of this partitioning changes over time: for some localities ceratopsian dinosaurs have higher carbon and oxygen isotope ratios than hadrosaurs, indicating a preference for plants living in open settings near the coast, whereas for other localities isotope ratios are lower, indicating a preference for plants in the understory of forests. In most cases the isotope ratios among hadrosaurs are similar and are interpreted to represent a dietary preference for plants of the forest canopy. The inferred differences in ceratopsian behavior are suggested to represent a change in vegetation cover and hence habitat availability in response to sea level change or to the position of river distributaries. Given our current lack of taxonomic resolution, it is not possible to determine if dietary and habitat preferences inferred from stable isotope data are associated with single, or multiple, species of hadrosaurian/ceratopsian dinosaurs.

The frequency of sublethal damage was examined in two ophiuroid species, Palaeocoma milleri and ?Ophioderma tenuibrachiata (Ophiodermatidae), from a shallow-water Jurassic “starfish bed.” None of 60 specimens was regenerating arms, a result that agrees with four previous studies of Paleozoic and early Mesozoic ophiuroids. By contrast, 66.1 percent of a living Ophioderma brevispinum population from Belize were regenerating one or more arms. For living populations of Ophiothrix oerstedi, the natural injury frequency was high at sites where the field mortality of tethered individuals was high. All predators that have been observed feeding on ophiuroids cause sublethal injuries, which would appear as regenerating arms in the fossil record. These results support the hypothesis that predation pressure on ancient ophiuroid populations was low and increased after the Jurassic.

Developmental constraints presumably operate by influencing patterns of variability: when development causes some features to vary more than others and when the level of variability is correlated with evolutionary change, then development can be said to constrain evolution. This idea was tested by examining the relationship between tooth variation and three other factors: developmental processes, tooth function, and evolutionary change. Data came from two lineages of viverravid carnivorans (Viverravidae, Carnivora) from the Paleogene of North America. Variability in cusp position was significantly correlated with position in the developmental cascade, with the amount of intercusp growth (when growth is relatively greater in some cusps than others), and with amount of evolutionary change. This indicates that tooth development exerts a local constraint on phenotypic variability and on the evolutionary response to functional selection, but comparative data suggest that the developmental constraint itself may evolve. Intense directional or stabilizing selection may modify the developmental cascade so that the constraint is either removed or modified to permit new evolutionary patterns. Thus development does not constrain evolution in an absolute sense, but rather introduces modifiable patterns of covariance among crown features. Both development and function seem to play important, intertwined roles in coordinating evolutionary change in mammalian molars.

The subfamily Equinae in the Great Plains region of North America underwent a dramatic radiation and subsequent decline as climate changed from warm and humid in the middle Miocene to cooler and more arid conditions during the late Miocene. Here we use ecological niche modeling (ENM), specifically the GARP (Genetic Algorithm using Rule-set Prediction) modeling system, to reconstruct the geographic distribution of individual species during two time slices from the middle Miocene through early Pliocene. This method combines known species occurrence points with environmental parameters inferred from sedimentological variables to model each species' fundamental niche. The geographic range of each species is then predicted to occupy the geographic area within the study region wherever the set of environmental parameters that constrain the fundamental niche occurs. We analyze changes in the predicted distributions of individual species between time slices in relation to Miocene/Pliocene climate change. Specifically, we examine and compare distribution patterns for two time slices that span the period from the mid-Miocene (Barstovian) Climatic Optimum into the early Pliocene (Blancan) to determine whether habitat fragmentation led to speciation within the clade and whether species survival was related to geographic range size. Patchy geographic distributions were more common in the middle Miocene when speciation rates were high. During the late Miocene, when speciation rates were lower, continuous geographic ranges were more common. Equid species tracked their preferred habitat within the Great Plains region as well as regionally throughout North America. Species with larger predicted ranges preferentially survived the initial cooling event better than species with small geographic ranges. As climate continued to deteriorate in the late Miocene, however, range size became irrelevant to survival, and extinction rates increased for species of all range sizes. This is the first use of ENM and GARP in the continental fossil record. This powerful quantitative biogeographic method offers great promise for studies of other taxa and geologic intervals.

The biogeography of differences in average shape of the modern planktonic foraminifer Globorotalia truncatulinoides (d'Orbigny) exhibits systematic relationships to changes in the ocean's surface environment. Comparison of these shape changes, as they exist today in the Southern Hemisphere, with fossil shapes preserved in a Late Pleistocene record from the South Atlantic Ocean, shows that the biogeography of G. truncatulinoides ecophenotypes has changed markedly through time. Beginning at least 700,000 yr ago and continuing up to the present, there has been a gradual but clear migration of certain morphotypes of G. truncatulinoides toward lower latitudes. The history of this migration bears no simple relationship to the cyclic climatic changes that characterize the Late Pleistocene. We conclude that either (1) phenotypic variants of Gr. truncatulinoides reflect some previously unmeasured, gradually changing aspect of Late Pleistocene oceans, or (2) we are witnessing a gradual evolution of the environment preferences of G. truncatulinoides.

An extreme, dorsally hyperextended posture of the spine (opisthotonus), characterized by the skull and neck recurved over the back, and with strong extension of the tail, is observed in many well-preserved, articulated amniote skeletons (birds and other dinosaurs, pterosaurs, and at least placental mammals). Postmortem water transport may explain some cases of spinal curvature in fossil tetrapods, but we show how these can be distinguished from causes of the opisthotonic posture, which is a biotic syndrome. Traditional biotic explanations nearly all involve postmortem causes, and have included rigor mortis, desiccation, and contraction of tendons and ligaments. However, examination of the process of rigor mortis and experimental observations of drying and salinity in carcasses of extant animals show that these explanations of the “dead bird” (opisthotonic) posture account for few or no cases. Differential contraction of cervical ligaments after death also does not produce the opisthotonic posture. It is not postmortem contraction but perimortem muscle spasms resulting from various afflictions of the central nervous system that cause these extreme postures. That is, the opisthotonic posture is the result of “death throes,” not postmortem processes, and individuals so afflicted assumed the posture before death, not afterward. The clinical literature has long recognized that such afflicted individuals perish from asphyxiation, lack of nourishment or essential nutrients, environmental toxins, or viral infections, among other causes. Accepting the actual causes of the opisthotonic posture as perimortem and not postmortem provides insights into the causes of death of fossilized specimens, and also revises interpretations of paleoenvironmental conditions of many fossil deposits. The opisthotonic posture tells us more about the circumstances surrounding death than about what happened after death. Finally, the opisthotonic posture appears to have a phylogenetic signal: it is so far reported entirely in ornithodiran archosaurs (dinosaurs and pterosaurs) and in crown-group placentals, though the distribution in mammals may expand with further study. It seems important that the opisthotonic posture has been observed extensively only in clades of animals that are known or thought to have high basal metabolic rates: hypoxia and related diseases would be most likely to affect animals with high oxygen use rates.

Convincing hypotheses of the origin of major invertebrate groups are difficult to make in the absence of phylogenetic analyses. In spite of this, several scenarios exist for the origin of the unusual echinoid order Clypeasteroida. I expand upon the most probable of these models by performing a phylogenetic analysis on three clypeasteroid suborders, the enigmatic fossil genus Togocyamus, and the extinct Oligopygoida. This analysis shows that the oligopygoids are the sister group of the Clypeasteroida plus Togocyamus. The latter is here considered a plesion (extinct sister group) to the crown group Clypeasteroida. Within that order, the suborder Clypeasterina is the sister group to the Laganina plus Scutellina. A new classification of all these taxa is presented. The phylogeny is based on 47 characters and incorporates data on external appendages, Aristotle's lantern anatomy, and test structure of irregular echinoids, as well as new information on the morphology of Togocyamus. The earliest clypeasteroids had a lantern similar to that of adult oligopygoids, which in turn inherited their lantern from a cassiduloid-like ancestor that retained the lantern into adulthood. This lantern is absent in adult cassiduloids. Subsequent changes, including modification of the lantern into a crushing mill, extreme flattening of the test, and proliferation of food-gathering tube feet have allowed clypeasteroids to become epifaunal inhabitants of environments characterized by fine, shifting substrates, a habitat previously inaccessible to most other irregular echinoids.

We calculated water vapor conductance (a product of eggshell porosity) from the first definitively identified sauropod egg (Megaloolithus patagonicus) from the Auca Mahuevo locality in Argentina. We then compared the results with those from M. siruguei (an egg type long associated with sauropod dinosaurs) from the Pinyes locality in Spain. The 14-cm Auca Mahuevo egg has a thinner eggshell and 47 times fewer pores than the 22-cm M. siruguei specimen. The resulting water vapor conductance (GH2O) of the titanosaur and M. siruguei eggs is 341 and 3979 mg H2O day−1 Torr−1, respectively; these values are two and ten times greater than in avian eggs of comparable size, but lower than in eggs of most modern reptiles. Clutches from Auca Mahuevo typically contain 20–40 eggs; in contrast, M. siruguei clutches from the Pinyes site average nine eggs. The GH2O of M. siruguei exceeds that of the Argentine egg by an order of magnitude, supporting previous inferences of egg burial. The GH2O of the Argentine titanosaur egg closely approximates that of Troodon and some oviraptorid eggs, previously calculated as equal to or two times greater than, respectively, the GH2O of avian eggs of similar size. Higher embryonic growth rates (relative to modern reptiles), especially in some dinosaurs with large clutch mass, may have required incubation in a more open environment, where water conservation represented a more critical factor than in a buried clutch. The lower GH2O calculated for the two megaloolithid eggs is consistent with previous interpretations of nesting mode that are based on site taphonomy and nesting traces. This study indicates that at least some dinosaurs did not fully bury their eggs.

The Paratethyan basins of eastern Europe and western Asia became isolated from marine influence in the Late Miocene, and were the sites of several remarkable endemic radiations of brackish and freshwater organisms. Here I describe the patterns of tempo and mode before and during the radiation of the gastropod Melanopsis in the Pannonian basin of eastern and central Europe, and I explore the underlying mechanisms of evolutionary change.

The most ancient melanopsid species in this area, M. impressa, was present in freshwater areas marginal to the basin well before the radiation. Widely spaced samples of M. impressa indicate that this species underwent a period of stasis lasting at least 7 m.y. The end of stasis corresponded with the extinction of the last of the normal marine fauna in the basin, suggesting that the lack of other fauna and/or reduced salinity in the basin permitted expansion of the melanopsids from the basin margins into the basin proper. Stasis ended with the onset of changes in size, shouldering, and ontogeny, which led eventually to M. fossilis. Change occurred over a 2-m.y. interval; a series of intermediates is present for all three characters. Within-sample correlations provide no evidence that the three characters are constructionally linked; instead they appear to be changing independently. The mode of change in the M. impressa–M. fossilis lineage appears to have been anagenetic. Alterations in the rate and direction of selection (and/or genetic links between characters) are probably required to explain the overall slowness of the change.

Most Pannonian basin melanopsid species arose by rapid cladogenesis in the Middle Pannonian Stage. Physical factors in the basin probably influenced the timing of this diversification; contrasting patterns of variation and diversity between two melanopsid clades suggest that intrinsic factors influenced the extent of diversification.

Bedding-plane assemblages of Ediacaran fossils from Mistaken Point, Newfoundland, are among the oldest known records of complex multicellular life on Earth (dated to ~565 Ma). The in situ preservation of these sessile but otherwise deeply enigmatic organisms means that statistical analyses of specimen positions can be used to illuminate their underlying ecological dynamics, including the interactions between taxa.

Fossil assemblages on Mistaken Point D and E surfaces were mapped to millimeter accuracy using differentiated GPS. Spatial correlations between 10 well-defined taxa (Bradgatia, Charniid, Charniodiscus, Fractofusus, Ivesheadiomorphs, Lobate Discs, Pectinifrons, Plumeropriscum, Hiemalora, and Thectardis) were identified using Bayesian network inference (BNI), and then described and analyzed using spatial point-process analysis. BNI found that the E-surface community had a complex web of interactions and associations between taxa, with all but one taxon (Thectardis) interacting with at least one other. The unique spatial distribution of Thectardis supports previous, morphology-based arguments for its fundamentally distinct nature. BNI revealed that the D-surface community showed no interspecific interactions or associations, a pattern consistent with a homogeneous environment.

On the E surface, all six of the abundant taxonomic groups (Fractofusus, Bradgatia, Charniid, Charniodiscus, Thectardis, and Plumeropriscum) were found to have a unique set of interactions with other taxa, reflecting a broad range of underlying ecological responses. Four instances of habitat associations were detected between taxa, of which two (Charniodiscus–Plumeropriscum and Plumeropriscum–Fractofusus) led to weak competition for resources. One case of preemptive competition between Charniid and Lobate Discs was detected. There were no instances of interspecific facilitation. Ivesheadiomorph interactions mirror those of Fractofusus and Charniodiscus, identifying them as a form-taxonomic grouping of degradationally homogenized taphomorphs. The absence of increased fossil abundance in proximity to these taphomorphs argues against scavenging or saprophytic behaviors dominating the E-surface community.

The published fossil record has significant bias in favor of common and biostratigraphically important taxa when compared with data obtained from a thorough examination of several hundred collections from the Western Interior of North America. Overall species diversity is underestimated by a factor of 3 to 4, and bivalve and gastropod diversity by a factor of 5. The proportion of bivalves increased from 40 to 56% of the fauna, and the proportion of ammonites decreased from 28 to 18%. Thirteen published reports listed 65 species from 203 reported occurrences. Data from all sources showed 170 species for 1050 occurrences. By using abundance data and assuming a log-normal distribution, as many as 200 fossilizable mollusc species may have inhabited the Western Interior during the uppermost biozone of the Cenomanian. The importance of this study is that it quantifies the bias in the published fossil record relative to the potential fossil record for an unusually well studied interval of geologic time. The bias would be greater for less well studied strata.

This paper presents a method for constraining the age of a clade with the ages of the earliest fossil specimens in that clade's outgroups. Given a sufficiently deep, robust, well-resolved, and stratigraphically consistent cladogram, this method can yield useful age constraints even in the absence of specific information about the fossil preservation and recovery rates of individual taxa. The algorithm is applied to simulated data sets to demonstrate that this method can yield robust constraints of clade ages if there are sufficient fossil outgroups available and if there is a finite chance that additional outgroups may be discovered in the future. Finally, the technique is applied to actual fossil data to explore the origin of modern placental mammals. Using data from recently published cladograms, this method indicates that if all Mesozoic eutherians are regarded as outgroups of Placentalia, then the last common ancestor of modern placental mammals and their Cenozoic allies lived between 65 and 88–98 million years ago, depending on the assumed cladogram and the number of outgroups included in the analysis.

Both fossil preservation and sampling methods affect perceived patterns of biotic diversity. Artificial range truncations, for example, may lead to incongruences between apparent- and actual-diversity curves. Thus, a catastrophic extinction event may appear gradual. Recent advances in biostratigraphic-gap analysis provide models for the distribution of gap lengths between fossil occurrence horizons and provide methods to place confidence intervals on local taxon ranges and remove the biases caused by artificial range truncations. Confidence intervals for a set of local taxon ranges may then be evaluated collectively to test a hypothesis of co-extinction/co-emigration or co-origination/co-immigration. In the case of terminal Cretaceous ammonites from Seymour Island, range-chart data are compatible with an abrupt extinction event, although the test statistic is not minimized at the stratigraphic horizon that was suggested by Macellari (1986).