New mid-Cretaceous stable isotope (δ18O and δ13C) records of multiple planktonic foraminiferal species and coexisting coccoliths from Blake Nose (western North Atlantic) document a major depth-ecology reorganization of planktonic foraminifera. Across the Albian/Cenomanian boundary, deep-dwelling Praeglobotruncana stephani and Rotalipora globotruncanoides adapted to living at a shallower depth, while, at the same time, the population of surface-dwelling Paracostellagerina libyca declined. Subsequently, the opportunistic species Hedbergella delrioensis shifted to a deep environment, and the deep-dwelling forms Rotalipora montsalvensis and Rotalipora reicheli first appeared. The primary paleoenvironmental cause of the observed changes in planktonic adaptive strategies is uncertain, yet their coincidence with an earliest Cenomanian cooling trend reported elsewhere implicates the importance of reduced upper-ocean stratification. Although there has been an implicit assumption that the species-specific depth habitats of fossil planktonic foraminifera were invariant through time, planktonic paleoecology is a potential variable. Accordingly, the possibility of evolutionary changes in planktonic foraminiferal depth ecology should be a primary consideration (along with other environmental parameters) in paleoceanographic interpretations of foraminiferal stable isotope data.

Some statistical methods are described for the detection and estimation of variability in the rate of evolution, from biometrical data on single characters in fossil lineages. The methods were applied to some published examples of protistan and mammalian lineages, extending between 0.8 and 6.4 Myr in duration. Of six cases examined, only one failed to show evidence for significant variation in the rate of evolution, and only four deviated significantly from a random walk pattern, despite significant net evolutionary change in all cases but one.

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.

We review and synthesize multiple biotic and abiotic proxies for marine nutrient and food availability, primary productivity, and food quality (stoichiometry) and propose what their relationships may have been to macroevolutionary processes, especially speciation. This review confirms earlier suggestions that there has been an overall increase in marine primary productivity over the Phanerozoic, but indicates that the increase has been irregular and that present levels may not be the peak. We integrate these indicators into a new estimate of relative primary productivity in the global ocean through the Phanerozoic. We then combine multiple, frequently conflicting ecological-evolutionary hypotheses into a general model for how primary production may affect speciation over geological time scales. This model, an elaboration and extension of the “speciation cycle” previously proposed by Grant and Grant, attempts to explain why an increase in food supply sometimes is associated with decreased diversity, and at other times with increased diversification. We propose some simple tests for the application of this model to the fossil record.

Mathematical modeling of cladogenesis and fossil preservation is used to explore the expected behavior of commonly used measures of taxonomic diversity and taxonomic rates with respect to interval length, quality of preservation, position of interval in a stratigraphic succession, and taxonomic rates themselves. Particular attention is focused on the independent estimation of origination and extinction rates. Modeling supports intuitive and empirical arguments that single-interval taxa, being especially sensitive to variation in preservation and interval length, produce many undesirable distortions of the fossil record. It may generally be preferable to base diversity and rate measures on estimated numbers of taxa extant at single points in time rather than to adjust conventional interval-based measures by discarding single-interval taxa.

A combination of modeling and empirical analysis of fossil genera supports two major trends in marine animal evolution. (1) The Phanerozoic decline in taxonomic rates is unlikely to be an artifact of secular improvement in the quality of the fossil record, a point that has been argued before on different grounds. (2) The post-Paleozoic rise in diversity may be exaggerated by the essentially complete knowledge of the living fauna, but this bias is not the principal cause of the pattern. The pattern may partly reflect a secular increase in preservation nevertheless.

Apparent temporal variation in taxonomic rates can be produced artificially by variation in preservation rate. Some empirical arguments suggest, however, that much of the short-term variation in taxonomic rates observed in the fossil record is real. (1) For marine animals as a whole, the quality of the fossil record of a higher taxon is not a good predictor of its apparent variability in taxonomic rates. (2) For a sample data set covering a cross-section of higher taxa in the Ordovician, most of the apparent variation in origination and extinction rates is not statistically attributable to independently measured variation in preservation rates. (3) Previous work has shown that standardized sampling to remove effects of variable preservation and sampling yields abundant temporal variation in estimated taxonomic rates. While modeling suggests which rate measures are likely to be most accurate in principle, the question of how best to capture true variation in taxonomic rates remains open.

It is now widely recognized that a large number of Cretaceous planktic foraminiferal species are commonly found associated with fully Danian faunas in many K/T boundary sections and deep-sea cores. This “Cretaceous” fauna has traditionally been regarded as representing the reworking of older Cretaceous sediments into younger strata, though recent isotopic data from some species indicates that, at least in these instances, the reworking hypothesis is false. To further test this reworking hypothesis the biogeography of this “Cretaceous” fauna is compared to the underlying uppermost Maastrichtian biogeography and to the biogeography of lowermost Danian planktic foraminiferal faunas. Results show that there is no regular decline in species richness, extinction, or faunal co-occurrence values for this “Cretaceous” fauna at progressively higher (=younger) Danian stratigraphic horizons. Moreover, there is no compelling association between the stratigraphic persistence of this “Cretaceous” fauna and shallow depositional settings. Instead, this fauna is characterized by: (1) a close (and predictive) association between “Cretaceous” and indigenous Danian species richness values throughout the lower Danian, (2) a close numerical and geographic correspondence between Danian speciation and the disappearance of “Cretaceous” species from the Danian fossil record, and (3) a pronounced similarity between changes in the general biogeographic structures of the “Cretaceous” and associated Danian faunas throughout the study interval. These data suggest that the K/T planktic foraminiferal extinction event exhibited a marked geographic structure with low and middle latitude faunas experiencing differentially high extinction rates in the lowermost Danian zones P0 and P1a and high latitude survivor faunas persisting more or less unchanged into the overlying zone, P1b and P1c. Taken together, these results challenge the traditional concept of an instantaneous uppermost Cretaceous planktic foraminiferal mass extinction and its proposed causal connection to bolide impact.

Traditional wisdom of the evolution of lungs in fishes is that lungs arose when gill ventilation was hindered by an aquatic habitat that was low in oxygen. This scenario has been buttressed primarily by a proposed correlation between extant air-breathing fishes and hypoxic habitats, as well as by the fact that early vertebrate fossils were found in sediments believed to indicate a semi-arid environment. There are problems with this scenario, yet it retains a dominant influence on how the evolution of aerial respiration is viewed. This paper presents a new hypothesis for lung evolution that is more consistent with the fossil record and physiology of extant animals than the traditional scenario; I propose that lungs evolved to supply the heart with oxygen. The primitive vertebrate heart was spongy in architecture and devoid of coronary support, obtaining oxygen from luminal blood. By supplying oxygen to this tissue, lungs may have been important in ancient fishes for sustaining activity, regardless of environment. Furthermore, this function for lungs may have influenced cardiovascular adaptations of tetrapods because their divided cardiovascular system isolates the right side of the heart from pulmonary oxygen. I propose that three innovations compensate for this isolation: In extant amphibians oxygen-rich blood from cutaneous and buccal respiration enters the right side of the heart; in chelonians and lepidosaurs the intracardiac shunt washes oxygen-rich blood into the right side of the heart; in mammals, birds, and perhaps in crocodilians, support of the heart by coronary vasculature eliminates this problem.

A survey was undertaken of the developmental types of the neogastropod families Volutidae, Fasciolariidae, Nassariidae, Mitridae, Buccinidae and Olividae in the Palaeocene and Eocene series of the Gulf of Mexico. Although the families were dominated by planktotrophic development in the Paleocene, most show a general increase in numbers of species with nonplanktotrophic development through the Eocene. Present day environmental correlations with larval development do not adequately explain the observed shift in developmental types in the Early Tertiary. It is proposed that higher speciation rates for nonplanktotrophs and a unidirectional trend in developmental change may account for the progressive increase of nonplanktotrophic development.

Approximately half of the Caribbean Oligocene reef coral fauna became locally extinct during the Early Miocene; roughly two thirds of the genera driven to local extinction still survive in the Indo-Pacific. Coral genera with lecithotrophic larvae (brooders) preferentially survived, over those with planktotrophic larvae (broadcasters). Among 37 genera for which we inferred reproductive mode, 73% of brooding genera survived the Oligocene/Miocene extinction events, while only 29% of the broadcasting genera survived. The proportion of brooders to broadcasters also increased markedly. During the late Oligocene, 47% of Caribbean reef coral genera were broadcasters, but in the middle Miocene, only 32% of the genera were broadcasters.

Survivorship in Puerto Rican reefs was correlated with tolerance of cold and turbid conditions. Genera tolerant of both cold water and turbidity had much higher survival rates than those tolerant of turbidity alone. Only 25% of the genera that could tolerate neither cold water nor turbidity survived. Most of the eurytopic genera were brooders, while most of the stenotypic genera were broadcasters.

We present two hypotheses that may account for the preferential survivorship of brooders: the recruitment hypothesis, and the dispersal hypothesis. The recruitment hypothesis holds that brooders survive preferentially because lecithotrophic larvae have higher recruitment success than do planktotrophic larvae in marginal habitats, such as upwelling zones. This is supported by the correlation of brooding and eurytopy. The dispersal hypothesis suggests that brooders survive preferentially because lecithotrophic larvae, which typically inherit zooxanthellae from the egg, have a longer larval lifespan and, hence, a wider potential dispersal range, than planktotrophic larvae, which typically capture zooxanthellae from the water column. Biogeographic range data, however, do not support this second hypothesis: modern Indo-Pacific brooding and broadcasting genera have nearly identical ranges, and many brooding species have narrower longitudinal ranges than do broadcasting species. Preferential survivorship of brooding corals contrasts sharply with survivorship patterns among molluscs during extinction events; among molluscs, broadcasters are favored over brooders.

A major increase in upwelling at the Oligocene/Miocene boundary was probably responsible for this extinction/geographic restriction event. Preferential survival of brooding and mixed mode coral genera appears to be a product of their being better able to recruit and survive in marginal conditions such as upwelling zones.

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.

Geographic range size is one of the few traits that promoted survivorship during both mass and background extinctions, but the exact reason (or reasons) why a large geographic range confers extinction resistance remains unclear. Proposed explanations have focused on the roles of dispersal ability, climate tolerance, global abundance, and widespread ranges in predicting taxon longevities. This study uses biogeographic data for late Paleozoic brachiopod genera to test the relative contribution of these traits to genus longevities, using simple but accurate proxy measurements. The results demonstrate a strong positive relationship between genus longevity and geographic range size, which is robust to several potential errors. Further, latitudinal range, which predominantly reflects climate tolerance, was no more important than longitudinal range, which predominantly reflects dispersal ability, in predicting genus longevities. Rather, longevities were an outcome of the total number of occurrences, which estimates global abundance, and the advantages of widespread distribution, regardless of which particular traits were responsible for generating the total geographic range. The advantages of a large geographic range were apparent during both background and mass extinctions of late Paleozoic time. Although not statistically significant, there was a tendency for the greatest selectivity to occur in intervals with the lowest extinction rates. The correlation of genus longevity and geographic range size had a profound consequence for the secular pattern of global brachiopod diversity: because the diversity of genera with small geographic ranges was more volatile owing to their correspondingly shorter longevities, global diversity and mean geographic range size paralleled each other almost exactly. Given that the correlation between taxon longevity and geographic range size has also been demonstrated for other taxonomic groups and at other time intervals, these results suggest that global diversity curves compiled from taxonomic databases dominantly reflect changes in the diversity of genera with small geographic ranges.

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.

Two communities dominated by deposit-feeding bivalves are recognized in Quisset Harbor, Massachusetts. Interference with life position and sediment preconditioning restrict the distribution of some species. The Eel Grass Community, dominated by Nucula proxima, comprises primarily non-siphonate forms which live in fecal-pellet rich, high-water-content, oxygenated sediment. The sediment texture results from active burrowing. This soft, mobile sediment readily clogs the siphons of siphonate bivalves. The Channel Community, dominated by Yoldia limatula, comprises primarily siphonate forms which live in partly compacted, low-water-content muds. Each community is characterized by abundant surface, shallow, and deep feeding species. Competition for food has generated this distinct stratification in feeding depth of the dominant species with each community.

Similar competitive relationships, affecting both species occurrence and trophic stratification, are apparent in three Silurian age deposit-feeding bivalve communities from the McAdam Brook Formation of Nova Scotia. A community dominated by four non-siphonate species inhabited soupy, high-water-content muds. A community dominated by three siphonate species lived in cohesive muds. A mixed community, dominated by four species, two non-siphonate and two siphonate species (the most abundant of which are present, but not as the most abundant forms, in the other communities), characterized the mid-way point of the ecotone between the other two communities.

These deposit-feeding bivalve dominated communities have similar structural patterns although separated by 400 million years and despite environmental differences and evolutionary changes. Both response to physical factors, such as sediment character, and interspecific competition impose and maintain this type of community structure.

Systematics and paleontology have had a long conceptual relationship, united by the common goal of reconstructing the history of life. Yet, with few exceptions, paleontologists have had little input into formulating systematic theory and methodology. The reasons for this apparently relate to two conceptual-philosophical traditions of post-Darwinian paleontology: (1) the widespread adoption of a species concept in which taxa are viewed as nondiscrete, arbitrarily designated segments of evolutionary continua, and (2) the belief that phylogenetic reconstruction is primarily an empirical matter of tracing evolutionary change through the stratigraphic record.

Available systematic evidence supports the hypothesis that species are real, discrete units in space and time and that, unless they are postulated to be directly ancestral to another species, they can be defined by the possession of one or more evolutionary novelties (derived characters). Species beginnings are delineated by speciation (vicariance) events and their terminations by subsequent speciation events or by extinctions.

Natural groups are composed of taxa that have shared a common genealogical history. Cladistic analysis is a method to construct and test hypotheses of monophyly and thereby define natural groups. Cladistic hypotheses are necessary to investigate many of the major questions within contemporary paleobiology. Virtually no studies of evolutionary rates, patterns of taxonomic diversity, modes of taxic evolution, and patterns of morphological diversification can be undertaken without reference to cladistic hypotheses about the composition of natural groups.

Because paleobiology is historical in its content, paleontologists are greatly limited in their ability to use paleontological data to investigate questions about the evolutionary process. According to current evolutionary theory, the concepts of adaptation and natural selection relate to genetic and ecological processes that take place within local populations (microevolution). If so, then data relevant to examining these phenomena are likely to be lacking in paleontological samples. Consequently, explanations of paleontological pattern that include process-related concepts such as adaptation and natural selection are axiomatic in their logical structure and thus cannot be falsified or critically evaluated by that paleontological pattern.

We present a morphometric analysis of water transport cells within a physiologically explicit three-dimensional space. Previous work has shown that cell length, diameter, and pit resistance govern the hydraulic resistance of individual conducting cells; thus, we use these three parameters as axes for our morphospace. We compare living and extinct plants within this space to investigate how patterns of plant conductivity have changed over evolutionary time. Extinct coniferophytes fall within the range of living conifers, despite differences in tracheid-level anatomy. Living cycads, Ginkgo biloba, the Miocene fossil Ginkgo beckii, and extinct cycadeoids overlap with both conifers and vesselless angiosperms. Three Paleozoic seed plants, however, occur in a portion of the morphospace that no living seed plant occupies. Lyginopteris, Callistophyton, and, especially, Medullosa evolved tracheids with high conductivities similar to those of some vessel-bearing angiosperms. Such fossils indicate that extinct seed plants evolved a structural and functional diversity of xylem architectures broader, in some ways, than the range observable in living seed plants.

Forty-six specimens of Nautilus pompilius Linnaeus were captured in depths varying between 100 and 500 m outside of the fringing reef near Suva, Fiji Islands. Thirty-eight of the specimens were male. Air weight per individual varied between 347 and 630 g. Sexual dimorphism in size is indicated, since mature shell modifications (approximated septa, blackened aperture) were present in two females weighing about 350 g (soft parts plus shell) and one weighing slightly over 400 g; the smallest male showing mature shell modifications weighed 496 g. All newly captured specimens were heavier than seawater, with mean weight in seawater of 1.87 g determined for twenty-five specimens. Total volumes of cameral liquid ranged between 13.5 and 0 ml. Thirteen of twenty-five sampled specimens showed less than 1.0 ml of cameral liquid from all chambers. Average cameral liquid osmolarity was lower than that observed in sampled populations of N. macromphalus from New Caledonia and N. pompilius from the Philippine Islands. Maximum swimming rates were 0.25 m/sec. N. pompilius exhibits two common color polymorphs.

Ecological ordination can reveal gradients in the species composition of fossil assemblages that can be correlated with paleoenvironmental gradients. Ordinations of simulated data sets suggest that nonmetric multidimensional scaling (NMDS) generally produces less distorted results than detrended correspondence analysis (DCA). We ordinated 113 brachiopod-dominated samples from the Frasnian (Late Devonian) Brallier, Scherr, and lower Foreknobs Formations of southwest Virginia, which represent a range of siliciclastic marine paleoenvironments. A clear environmental signal in the ordination results was obscured by (apparently) opportunistic species that occurred at high abundance in multiple environments; samples dominated by these species aggregated in ordination space regardless of paleoenvironmental provenance. After the opportunist-dominated samples were removed, NMDS revealed a gradient in species composition that was highly correlated with substrate (grain size); a second, orthogonal gradient likely reflects variation in disturbance intensity or frequency within grain-size regimes. Additional environmental or ecological factors, such as oxygenation, may also be related to the gradients. These two gradients, plus the environmental factors that controlled the occurrence of opportunistic species, explain much of the variation in assemblage composition in the fauna. In general, the composition of fossil assemblages is probably influenced by multiple paleoecological and paleoenvironmental factors, but many of these can be decomposed and analyzed.

Species arise and establish themselves over the geologic time scale. This process is manifested as a change in the relative frequency of occurrences of a given species in the global pool of species. Our main goal here is to model this rise and the eventual decline of microfossil species using a mixed-effects model where groups each have a characteristic occurrence trajectory (main effects) and each species belonging to those groups is allowed to deviate from the given group trajectory (random effects). Our model can be described as a “hat” with logistic forms in the periods of increase and decline. Using the estimated timings of rises and falls, we find that the lengths of the periods of rise are about as long as the lengths of the periods when species are above 50% of their estimated maximal occurrences. These latter periods are here termed periods of dominance, which are in turn about the same length as the species' periods of fall. The peak rates of the rises of microfossils are in general faster than their peak rates of falls. These quantified observations may have broad macroevolutionary and macroecological implications. Further, we hypothesize that species that have experienced and survived high levels of environmental volatility (specifically, periods of greater than average variation in temperature and productivity) during their formative periods should have longer periods of dominance. This is because subsequent environmental variations should not drive them to decline with ease. We find that higher estimated environmental volatility early in the life of a species positively correlates with lengths of periods of dominance, given that a species survives the initial stress of the environmental fluctuations. However, we find no evidence that the steepness of the rise of a species is affected by environmental volatility in the early phases of its life.

Analysis of molluscan collections from a 3+ m.y. interval around the Cretaceous–Tertiary (K–T) interval in east Texas suggests that molluscs suffered an extinction at or near the K–T boundary, followed by a prolonged period of stress which lasted through the P0 and P1a planktic foraminiferal zones. The stressed period was characterized by low species richness, low abundances of individuals, high species turnover and a dominance of deposit feeders. Species richness and the relative abundance of deposit feeders generally track the 13C depletion curve suggesting that the stress was caused by a lack of primary production. A stable, relatively diverse, suspension feeding molluscan community was reestablished less than two million years after the K–T boundary. The total number of species within the habitat did not recover to Cretaceous levels within the study interval.

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.