Many studies have examined temporal changes in insect feeding on angiosperm leaves, but none have considered variability within a single stratigraphic level. If spatial variability within a level is high, a single sample will not adequately represent the level and may either mask true temporal changes or create spurious ones. In order to measure the spatial variability in fossil insect feeding damage, I collected 12 replicate samples from two laterally extensive carbonaceous shale beds (55.2 and 52.6 Ma) from the early Eocene of the Bighorn Basin, Wyoming. Over 2800 fossil angiosperm leaves were scored for presence or absence of 50 insect damage morphotypes. Damage frequency, diversity, and composition were computed for both the bulk flora and individual plant species in each sample, and variation within a bed was compared with differences between the two beds. Differences in diversity and composition between beds were significantly greater than variations within a bed, and intra-bed variation was primarily due to differing floral composition. Damage frequency within a bed, however, was more variable than diversity. Damage diversity and composition reflect the number of insect species present, whereas damage frequency also depends on the number of insects present, which may be much more variable over small distances.

Thalattosuchians are crocodylomorphs mainly known from marine strata of Early Jurassic to Early Cretaceous age. They represent the earliest crocodylomorph radiation to an aquatic habitat and their evolutionary history offers very few records from freshwater settings. Here, we report several exquisitely preserved thalattosuchian skulls attributed to a derived teleosaurid from a pedogenic horizon located at the base of a fluvial series of alternating silts and sandstones of the Phu Kradung Formation (Upper Jurassic) of northeastern Thailand. Using laser ablation multicollector inductively coupled mass spectrometry (MC-ICP-MS) on tooth enamel and dentine, we measured isotopic ratios of strontium (87Sr/86Sr) to test the habitat of these teleosaurids. In addition, Sr concentrations of the dental tissues were estimated from the calibrated signal intensities of the Sr isotope measurements. The dataset includes bioapatite (teeth or scales) of eight terrestrial and five aquatic vertebrates. Theropods exhibit lower Sr concentrations both in enamel and dentine compared to others groups, a pattern in accordance with the calcium biopurification process, which predicts that Sr concentrations in the body of vertebrates decrease up the trophic chain. It also excludes the possibility that diagenesis has completely overprinted the Sr isotope compositions of the fossil assemblage, which exhibits a homogeneous 87Sr/86Sr signature above the Late Jurassic seawater value. Values for teleosaurid teeth are in the range of other values for vertebrates in the continental assemblage and imply that these crocodylomorphs did not migrate between freshwater and marine habitats at least in the time constraint of the mineralizing tooth. This result represents the first demonstration that a population of teleosaurids was established for a prolonged time in a freshwater environment. Whether the ability of teleosaurids to inhabit freshwater habitats is a secondary adaptation or whether it is plesiomorphic and inherited from freshwater ancestors is discussed.

Niche conservatism is increasingly recognized in diverse modern ecological settings, and it influences many aspects of modern ecosystems, including speciation mechanisms, community structure, and response to climate change. Here, we investigate the stability of niches with benthic marine invertebrates along a Late Ordovician onshore-offshore gradient on the Cincinnati Arch in the eastern United States. Using a Gaussian niche model characterized by peak abundance, preferred environment, and environmental tolerance, with these parameters estimated through weighted averaging and logistic regression, we find evidence of strong niche conservatism in peak abundance and preferred environment, particularly for abundant taxa. This conservatism is maintained in successive depositional sequences and through the nearly 9–10 Myr study interval. Environmental tolerance shows no evidence of conservatism, although numerical simulations suggest that the error rates in estimates of this parameter are so high that they could overwhelm evidence of conservatism. These numerical simulations also indicate that both weighted averaging and logistic regression produce useful estimates of peak abundance and preferred environment, with slightly better results for weighted averaging. This evidence for niche conservatism suggests that long-term shifts of higher taxa of marine invertebrates into deeper water are primarily the result of differential rates of origination and extinction. These results also add to the evidence of long periods of relatively stable ecosystems despite regional environmental perturbations, and they constrain the causes of peaked patterns in occupancy.

Particle size reduction is a primary means of improving efficiency in herbivores. The mode of food particle size reduction is one of the main differences between herbivorous birds (gizzard) and mammals (teeth). For a quantitative comparison of the efficiency of food comminution, we investigated mean fecal particle sizes (MPS) in 14 herbivorous bird species and compared these with a data set of 111 non-ruminant herbivorous mammal species. In general MPS increased with body mass, but there was no significant difference between birds and mammals, suggesting a comparable efficiency of food processing by gizzards and chewing teeth. The results lead to the intriguing question of why gizzard systems have evolved comparatively rarely among amniote herbivores. Advantages linked to one of the two food comminution systems must, however, be sought in different effects other than size reduction itself. In paleoecological scenarios, the evolution of “dental batteries,” for example in ornithopod dinosaurs, should be considered an advantage compared to absence of mastication, but not compared to gizzard-based herbivory.

The late Mesozoic through early Cenozoic is an interval of significant biologic turnover and ecologic reorganization within marine assemblages, but the timing and causes of these changes remain poorly understood. Here, we quantify the pattern and timing of shifts in the diversity (richness and evenness) and ecology of local (i.e., sample level) mollusk-dominated assemblages during this critical interval using field-collected and published data sets from the U.S. Gulf Coastal Plain. We test whether the biologic and ecologic patterns observed primarily at the global level during this time are also expressed at the local level, and whether the end-Cretaceous (K/Pg) mass extinction and recovery moderated these trends. To explore whether environment had any effect on these patterns, we examine data from shallow subtidal and offshore settings.

Assemblages from both settings recovered to pre-extinction diversity levels rapidly, in less than 7 million years. Following initial recovery, diversity remained unchanged in both settings. The trajectory of ecological restructuring was distinct for each setting in the wake of the K/Pg extinction. In offshore assemblages, the abundance and number of predatory carnivorous taxa dramatically increased, and surficial sessile suspension feeders were replaced by more active suspension feeders. In contrast, shallow subtidal assemblages did not experience ecological reorganization following the K/Pg extinction. The distinct ecological patterns displayed in each environment follow onshore-offshore patterns of innovation, whereby evolutionary novelties first appear in onshore settings relative to offshore habitats. Increased predation pressure may explain the significant ecological restructuring of offshore assemblages, whereby the explosive radiation of predators drove changes in their prey. Habitat-specific ecological restructuring, and its occurrence solely during the recovery interval, implies that disturbance and incumbency were also key in mediating these ecological changes.

A range of specialized burrowing sculptures evolved convergently in at least one lingulid and in several obolid and lingulasmatid genera. The sculptures in Lingula punctata (Ordovician) indicate a burrowing process with the pedicle trailing behind the shell, similar to that of Recent Lingulidae. In the Obolidae and Lingulasmatidae, the orientation of the sculptures indicates a burrowing process with the pedicle oriented downward in the sediment. A burrowing mechanism in which the valves alternated in functioning as anchors, without active participation of the pedicle, is proposed for these forms. Infaunal life habits for several obolid genera are further indicated by the shell morphology and by the distribution of repaired shell damage.

The burrowing Obolidae are likely to have been adapted to relatively high-energy environments, which required frequent reburrowing in addition to escape (upward) burrowing. The increase in bioturbation in these environments in the Middle Paleozoic may have contributed to their extinction. The Lingulidae, instead, being mostly adapted to extreme environmental conditions, could survive in marginal niches which were not intensively exploited by actively burrowing organisms.

Teeth and other dermal skeletal elements from three osteolepid and two eusthenopterid rhipidistians were examined with the scanning electron microscope to establish whether the hypermineralized outer layer was enameloid or enamel. Using Smith's (1978) morphological criteria to infer the developmental history of enamel and enameloid, enamel was found in the teeth and dermal bones of the osteolepids, whereas enameloid was identified in scales. Hyneria and Eusthenopteron teeth also had enamel. As enamel has also been found in teeth of living lungfishes and extant Latimeria, the presence of enamel in tetrapod teeth represents a primitive character retained from a lobe-finned ancestor and is not useful for phylogenetic studies within the lobe-finned/tetrapod group. Shellis and Miles' (1974) hypothesis that the transition from enameloid to enamel occurred via a fairly simple regulatory change is discussed.

Analysis of dental wear and damage is becoming an increasingly important tool in unraveling the trophic ecology of a wide range of vertebrates, and when applied to fossils it can provide evidence of both diet and feeding kinematics that is independent of morphological analysis. Conodonts have the best fossil record among vertebrates and their skeletal elements are known to exhibit surface wear and damage generated in vivo as a consequence of their function as teeth. We report the results of the first systematic survey and analysis of the frequency and extent of this wear and damage in conodonts (based on P1 elements from a range of Carboniferous genera). This has revealed that wear and damage are remarkably common, present in all conodont elements sampled. Multivariate analysis reveals that patterns of wear and damage differ significantly among different conodont taxa, and exploratory ANOVA and linear discriminant analyses show that wear and damage differ according to the position of taxa in an onshore-offshore gradient, and whether they are likely to have had a benthic or pelagic mode of life. The incidence of denticle tip spalling in particular is higher in more-offshore environments and in taxa likely to have had a pelagic mode of life. Aspects of the data also reflect the occlusal kinematics of the elements, providing a means of testing hypotheses of element function. Our results have wide-ranging implications for unlocking the fossil record of conodonts, by, for example, furnishing direct evidence of the diet-mediated processes that may have driven observed patterns of evolutionary change, and reducing the confounding effects of depth segregation when using conodonts in isotope-based paleotemperature studies.

We show that the decline in the extinction rate during the Phanerozoic can be accurately described by a logarithmic fit to the cumulative total extinction. This implies that extinction intensity is falling off approximately as the reciprocal of time. We demonstrate that this observation alone is sufficient to explain the existence of the proposed power-law forms in the distribution of the sizes of extinction events and in the power spectrum of Phanerozoic extinction, results that previously have been explained by appealing to self-organized critical theories of evolutionary dynamics.

One testable hypothesis of the theory that dinosaurs were endothermic is the observation that sauropod dinosaurs were too large, their heads were too small, and their food was too indigestible for them to be warm-blooded. Calculations on the daily calorie requirements of the sauropod Brachiosaurus, adjusted for digestibility and the energetic cost of “free-living,” were compared with the caloric density of Late Jurassic food plants and the feeding rates of an elephant and a giraffe. Using Brachiosaurus as a model I concluded that endothermy in large sauropods (greater than 55 metric tons) was impossible. Depending on assumptions about feeding rates and the cost of free-living, endothermy in smaller sauropods ranges from improbable to impossible.

Morphologic analysis was made on early whorls of two Paleozoic and 43 Mesozoic ammonoid species to obtain basic data on early life history. A positive linear relationship between protoconch and ammonitella sizes is recognized among the species examined. Total rotation angle of the ammonitella (ammonitella angle) seems to be independent of ammonitella size but has a significant negative linear relation with whorl expansion rate. This supports the model that the embryonic shell of ammonoids consists of gas-filled protoconch and the succeeding body whorl up to the primary varix. Growth analysis of siphuncle diameter in the early post-embryonic stage also indicates that the shell growth rates of the species hatched from large-sized eggs were smaller than those of species hatched from small-sized eggs.

A developmental model, based upon murine rodents, has been proposed by Kavanagh et al. (2007) to explain lower molar proportions in mammals. We produce a clade-wide macroevolutionary test of the model using the dental evolutionary trends in a unique radiation of extinct mammals endemic to South America (“Meridiungulata”) that comprise a diverse array of molar morphologies. All of the South American ungulate groups examined follow the inhibitory cascade model with the exception of two groups: Interatheriidae (Notoungulata) and Astrapotheria. For most taxa studied, ratios between lower molar areas are greater than 1.0, indicating a weak inhibition by m1 on the subsequent molars in the tooth row, and a trend to greater absolute size of the posterior molars. Comparisons of mean ratios between clades indicate that a significant phylogenetic signal can be detected, particularly between the two groups within Notoungulata— Typotheria and Toxodontia. Body mass estimates were found to be significantly correlated with both m3/m1 and m2/m1 ratios, suggesting that the larger body size achieved the weaker inhibition between the lower molars. Molar ratio patterns are examined and discussed in relation to the independent and numerous acquisitions of hypsodonty that are characteristic of dental evolution in “Meridiungulata.”

The temporal dimension of fossil sequences provides a critical component to the study of intraspecific dynamics and species formation. Here I report on the branching and subsequent morphological evolution of two gastropods (Melanopsis fossilis and M. vindobonensis) from the Late Miocene of the Pannonian basin in eastern and central Europe. Although morphological divergence between species is rapid, intermediates between the two species co-occur with typical individuals for approximately 1 m.y. and then disappear. The long-term persistence of intermediates followed by their ultimate disappearance is a pattern that, to my knowledge, has not been previously observed.

Distinguishing genetic from ecophenotypic influences on shell form in freshwater prosobranchs is difficult. Nevertheless, consideration of the temporal, geographic, lithologic, and paleoecologic patterns of this sequence suggests that the morphologic differences between M. fossilis and M. vindobonensis had some genetic basis. Whether these forms were initially morphs of a single species or two species with some hybridization between them is impossible to determine. In either case, the morphological changes that resulted in M. vindobonensis were rapid, but the attainment of complete isolation between M. fossilis and M. vindobonensis apparently did not occur until approximately 1 m.y. later.

Scleractinian corals have two fundamentally different life strategies, which can be inferred from morphological criteria in fossil material. In the non-photosymbiotic group nutrition comes exclusively from heterotrophic feeding, whereas the photosymbiotic group achieves a good part of its nutrition from algae hosted in the coral’s tissue. These ecologic differences arose early in the evolutionary history of corals but with repeated evolutionary losses and presumably also gains of symbiosis since then. We assessed the biodiversity dynamics and environmental occupancy of both ecologic groups to identify times when the evolutionary losses of symbiosis as inferred from molecular analyses might have occurred and if these can be linked to environmental change. Two episodes are likely: The first was in the mid-Cretaceous when non-symbiotic corals experienced an origination pulse and started to become more common in deeper, non-reef habitats and on siliciclastic substrates initiating a long-term offshore trend in occupancy. The second was around the Cretaceous/Paleogene boundary with another origination pulse and increased occupancy of deep-water settings in the non-symbiotic group. Environmental factors such as rapid global warming associated with mid-Cretaceous anoxic events and increased nutrient concentrations in Late Cretaceous–Cenozoic deeper waters are plausible mechanisms for the shift. Turnover rates and durations are not significantly different between the two ecologic groups when compared over the entire history of scleractinians. However, the deep-water shift of non-symbiotic corals was accompanied by reduced extinction rates, supporting the view that environmental occupancy is a prominent driver of evolutionary rates.

Geochemical tools, including the analysis of stable isotopes from fossil mammals, are often used to infer regional climatic and environmental differences. We have further developed an oxygen isotope aridity index and used oxygen (δ18O) isotope values and carbon (δ13C) isotope values to assess regional climatic differences between the southeastern and southwestern United States during the Pleistocene. Using data collected from previously published studies, we assigned taxa to evaporation-sensitivity categories by quantifying the frequency and magnitude of aridity index values (i.e., an average taxon δ18O value minus a site specific proboscidean δ18O value). Antilocapridae, Camelidae, Equidae, and Cervidae were identified as evaporation-sensitive families, meaning that a majority of their water comes from the food they eat, thus indicating that they are more likely to capture changing climatic conditions. Bovidae, Tayassuidae, and Tapiridae were identified as less sensitive families, possibly because of increased or more variable drinking behavior. While it is difficult to tease out individual influences on δ18O values in tooth enamel, the use of an aridity index will provide a more in-depth look at relative aridity in the fossil record. Greater aridity index values in the Southwest suggest a drier climate than in the Southeast during the Pleistocene, and δ13C values suggest that diet does not determine evaporation sensitivity. The combination of more-positive δ13C values and the lack of forest indicator taxa in the Southwest suggest that landscapes were more open than in the Southeast. Inferred higher aridity in the Southwest may indicate that aridity or seasonal aridity/precipitation, not temperature or pCO2, was a greater driver of C4 abundance during the Pleistocene. Collectively, these data suggest that regional climatic and environmental interpretations can be improved by using an aridity index and a more detailed understanding of mammalian paleobiology.

Larger body size has long been assumed to correlate with greater risk of extinction, helping to shape body-size distributions across the tree of life, but a lack of comprehensive size data for fossil taxa has left this hypothesis untested for most higher taxa across the vast majority of evolutionary time. Here we assess the relationship between body size and extinction using a data set comprising the body sizes, stratigraphic ranges, and occurrence patterns of 9408 genera of fossil marine animals spanning eight Linnaean classes across the past 485 Myr. We find that preferential extinction of smaller-bodied genera within classes is substantially more common than expected due to chance and that there is little evidence for preferential extinction of larger-bodied genera. Using a capture–mark–recapture analysis, we find that this size bias of extinction persists even after accounting for a pervasive bias against the sampling of smaller-bodied genera within classes. The size bias in extinction also persists after including geographic range as an additional predictor of extinction, indicating that correlation between body size and geographic range does not provide a simple explanation for the association between size and extinction. Regardless of the underlying causes, the preferential extinction of smaller-bodied genera across many higher taxa and most of geologic time indicates that the selective loss of large-bodied animals is the exception, rather than the rule, in the evolution of marine animals.

The statistical significance of differences in evolutionary rate between major taxonomic groups is evaluated using conventional chi-square techniques on stratigraphic range data. Romer's (1966) compilation of stratigraphic ranges of fossil mammals is used to determine whether orders differ significantly from each other in generic origination and extinction rates. The evolutionary histories of 2180 genera (primarily Cenozoic in age) are analyzed. Chi-square testing shows that significantly high or low (P ≥ 0.99) extinction or origination occurs in 15% of the testable cases. Significantly high or low evolutionary turnover in a taxon (orders in this case) we term taxotely. Significantly high turnover rate is equivalent to Simpson's tachytely and significantly low turnover is equivalent to his bradytely.

In the mammal data set, taxotely is largely attributable to the influence of South American endemics. Some of the effect is interpreted as an artifact of biases in the fossil record (or its study) and some is attributed to real biological aspects of mammalian evolution.

Recent phylogenetic studies of Paleozoic gastropods show the classification provided by the Treatise on Invertebrate Paleontology to include numerous highly polyphyletic taxa. Here we test whether this classification reflects limits on the range of possible designs, general architectural constraints, or common functional solutions for Paleozoic gastropods. Our test evaluates whether superfamilial/subordinal level archetypes as defined by the Treatise evolved more frequently than expected among 626 Late Cambrian–Middle Devonian species. Using a previously established phylogeny and five general shell features (spire angle, exhalent current position, base angle, umbilical width, and apertural inclination) we show that there are fewer gastropod morphotypes than expected given the frequency of change in these features. This is true even after accounting for architectural constraints implied by the data. Moreover, the most common morphotypes include significantly more species and evolved significantly more times than expected. These results imply a set of architectural attractors for lower–middle Paleozoic gastropods, consistent with ecomorphological theory that particular morphotypes are best suited for particular lifestyles. Thus, the Treatise classifications likely reflect these ecomorphological patterns within subclades rather than phylogenetic patterns.

Ciliary transport of particulate suspensions in water was studied in the mantles of the inarticulate brachiopod Lingula anatina, the articulate brachiopod Coptothyris grayi and in the ciliated epithelium of the scleractinian coral Montipora foliosa. In both brachiopod species a close correlation was found between the water currents induced by the lophophore and by the mantle. Lophophore and mantle are considered here as functionally integrated organs which are active in the transport and capture of food, accumulation and the removal of (pseudo-)faeces, the sorting of particles, the pumping of water and respiration.

The tuberculate surface of Montipora foliosa is lined with ciliated epithelium; the polyps are situated in the depressions of this surface. The particulate suspension within the labyrinth is constantly diluted: particles are funneled through the furrows but end up at the tips of the tubercles. On their way they pick up some mucus. While they concentrate on the tubercles they coalesce to form larger boluses that are easily removed. The tubercular tips thus act as “bolus accumulators” and the unciliated margins of the brachiopod mantles appear to have the same function. In this accumulated form the particles are probably prevented from being recycled through the food-capturing system.

Based on these observations a preliminary functional interpretation is presented of some fossil brachiopods: the Devonian rhynchonellids Uncinulus and Hypothyridina and the Strophomenida, with the genus Marginifera serving as an example of the latter order. The morphology of the inner shell surface, and the inferred configuration of the lining ciliated mantle, was modified in these taxa by a variety of structures such as spines, tubercles, diaphragms and trails. It is suggested that the mantle of the evolved Uncinulidae and of the Strophomenida is particularly suited for the dilution of particulate suspensions before they reach the lophophore and for the accumulation and removal of mucous pseudofaeces. The inhibitory effect of the constrictions on the circulation of water could be counteracted by local ciliary pumping. In the strophomenids the mantle became a specialized and elaborate organ, while the lophophore probably remained relatively simple. In the other articulate stocks the lophophore became the central food-gathering and pumping device and in general the morphology of the mantle was maintained simple.

Analysis of stratigraphically separated collections of Late Ordovician fossilized marine invertebrates suggests that our understanding of faunal changes through time may be enhanced by considering them in terms of a successional framework. The faunal succession in the Nicolet River Valley, St. Lawrence Lowlands, Quebec, began with the initial colonization of an offshore barren mud by two or three presumably opportunistic deposit-feeding species. Diversity, both of species and of trophic types, increased gradually, peaked, and then decreased slightly with some faunal shifts, especially in the numerical predominance of the epifaunal element. A three-fold repetition of this quiet-water succession seems in each case to have been terminated by an external physical perturbation. There also appears to be a second, separate successional sequence which became localized on a somewhat coarser-grained and perhaps less stable substratum as the paleoshoreline was approached, due to sediment filling of the basin.