The late Maastrichtian sediments of the Mullinax-1 and Mullinax-3 boreholes from Brazos River, Texas, offer pristine material. These cores are prime candidates for providing an extraordinary window into the ecology of Guembelitria, a key genus in the K/Pg mass extinction event, as well as information on the habitats of other neritic species. Stable oxygen and carbon isotope analyses were performed on six planktic species (Guembelitria cretacea, Globigerinelloides asper, Heterohelix globulosa, Paraspiroplecta navarroensis, Pseudoguembelina costulata, Rugoglobigerina rugosa) and three benthic genera (Gavelinella, Cibicides, and Lenticulina). Our records support the contention that Guembelitria was fully planktic, as indicated by its δ18O values, which overlap the other planktic species, despite its possible origin from a tychopelagic benthic ancestor. However, Guembelitria is distinctly ranked very low in δ13C values, which overlap the benthic records. The anomalously low δ13C values of Guembelitria may represent an isotopic disequilibrium due to fast shell growth, like in its modern analogue Gallitellia vivans. Another explanation may be that these values are attributable to a neustonic life mode in the uppermost part of the oceans, where photosynthesis is inhibited by high UV and the near absence of nutrients. Because these waters are not photosynthetically depleted, calcification using carbon directly from these waters should yield δ13C values consistent with those found in Guembelitria. The ecological strategy that Guembelitria species used to deal with the nutrient-poor surface-water environments was an opportunistic blooming during stressful times of Maastrichtian global warming events and later during the K-Pg catastrophe.

Using queueing theory and the fast Fourier transform, we develop a new approach to analyzing the diversity paths of fossil taxa. The methods employed focus on calculating the probabilities, for each taxonomic group, of arriving at particular values of the diversity at particular times, assuming that diversity paths are random walks. These probabilities, which are represented in probability matrices, allow us to establish a number of null hypotheses that can be used to test for randomness in the diversity paths of the taxonomic groups themselves. As the model employed here is conceptually similar to that employed by Gilinsky and Bambach (1986) in bootstrapping, one consequence of the approach is, in effect, an analytic solution of the bootstrap. A comparison of bootstrapping and analytic results in 15 taxa of marine invertebrates gives nearly identical results. The analytic approach is more general, however, and allows for the calculation of probability matrices under other model assumptions not treated here. We have also applied the probability matrices to a series of tests for nonrandomness as a whole, in contrast to the more usual method of testing particular clade statistics. The familial diversity paths of all 36 extinct orders of marine invertebrates with ten or more families were analyzed. In no case was the null hypothesis of randomness rejected at the 0.05 level. Given the model, and assuming that extinct taxa are representative of all taxa, the familial diversity paths of marine orders appear to behave as random walks.

Life-sized scale models and a flume were used to observe simulated feeding and hydrodynamics of the Paleozoic camerate crinoid, Pterotocrinus. Three orientations of the crown and filtration fan (downcurrent, upcurrent, upright) in a gentle (3.5 cm/sec) current were used. Maximum baffling of the current and eddying of water into feeding areas occurred where models were oriented like most living stalked crinoids, with a parabolic filtration fan perpendicular to the current and ambulacral side downcurrent. Facing the ambulacral side of the fan upcurrent reduced time available for entrapment of potential food particles. Calyx shape was important in modifying the nature of the filtration fan baffle. Overlapping (36%) of the pinnulate arms increased the baffling effect of the filtration fan. Conspicuous tegminal structures, the Pterotocrinus wing plates, only had a baffling effect where the fan was in an upright position or the arms were infolded between the wing plates. Species of Pterotocrinus possessing blade-like wing plates could have fed effectively in several different orientations. Evolutionary success of Pterotocrinus species with blade-like wing plates probably resulted from dual feeding and protective functions of the wing plates.

Genera of montane conifers first appeared in the fossil record in large numbers in the Middle Eocene (47-45 Ma) as widespread volcanism in the Cordilleran region provided scattered, cold upland tracts suited for them. Higher subalpine environments, generally above 2,000 m, probably were confined then to upper slopes of scattered strato-volcanos and basement highs covered with thick pyroclastic deposits. As relief increased in the middle Tertiary, and especially during the Pliocene-Quaternary, the area suited for a subalpine forest zone increased significantly, enabling montane conifers to spread into that higher, colder environment and form a new major forest zone. During glaciations the subalpine zone was much larger and probably from 900-1,000 m lower than at present. Spreading post-glacial warmth and drought account for its retreat to higher levels and for the discontinuous distribution of the forest and its species today. Recency of the subalpine adaptive zone is implied also by the close relations of its varieties and subspecies of herbaceous perennials, a few annuals, and pines to those in vegetation zones below it.

Intraspecific aggression, or agonism, is a widespread intrasexual selective behavior important to understanding animal behavioral ecology and reproductive systems. Such behavior can be studied either by direct observation or inferred from wound/scar frequency in extant species but is difficult to document in extinct taxa, limiting understanding of its evolution. Among extant archosaurs, crocodylians display extensive intrasexual aggression, whereas birds show extreme visual/vocal intersexual display. The evolutionary origin of this behavioral divergence, and pattern in non-avian dinosaurs, is unknown. Here we document the morphology, frequency, and ontogeny of intraspecific facial bite lesions (324 lesions) in a large sample of tyrannosaurids (202 specimens, 528 elements) to infer patterns of intraspecific aggression in non-avian theropods. Facial scars are consistent in position and orientation across tyrannosaurid species, suggesting bites were inflicted due to repeated/postured behavior. Facial scars are absent in young tyrannosaurids, first appear in immature animals (~50% adult skull length), are present in ~60% of the adult-sized specimens, and show aggressor:victim size isometry. The ontogenetic distribution of bite scars suggests agonistic behavior is associated with the onset of sexual maturity, and scar presence in approximately half the specimens may relate to a sexual pattern. Considered in a phylogenetic context, intraspecific bite marks are consistent and widely distributed in fossil and extant crocodyliforms and non-maniraptoriform theropods, suggesting a potential plesiomorphic behavior in archosaurs. Their absence in maniraptoriform theropods, including birds, may reflect a transition from boney cranial ornamentation and crocodylian-like intrasexual aggression to avian-like intersexual display with the evolution of pennaceous feathers.

The Corbulidae are one of a handful of a primarily marine bivalve clades that exhibit a remarkable radiation, marked by increased species richness and divergent morphologies, within a long-lived lake. For corbulids, this diversification occurred within the lower to middle Miocene Pebas Formation of western Amazonia. Only one taxon associated with this radiation (Anticorbula) remains extant. We conducted a series of phylogenetic analyses to characterize diversification of Corbulidae within the Pebas Formation and relate that diversification to geologically older freshwater corbulids from the Paleocene Fort Union Formation of the northern Great Plains (United States). We used these results, as well as a quantitative examination of morphospace occupation, to infer whether Pebasian corbulids represent a true species flock, and whether the lacustrine system represented by the Pebas Formation represents a cradle of, or reservoir for, freshwater corbulid diversity. We conducted two sets of phylogenetic analyses using shell morphology characters. A genus-level data set incorporated type species of freshwater corbulid genera, any Paleocene representatives of these genera, and selected brackish and marine corbulid genera. A species-level analysis added all described freshwater corbulid taxa to the genus-level matrix. Our results were highly resolved (few most-parsimonious trees), but not particularly robust (low branch support). For the genus-level matrix, we used a taxon jackknife procedure to explore the effects of taxon sampling on tree stability and topology. Jackknife results recover a subclade of freshwater taxa (including both Anticorbula and Pachydon species and the Paleocene Ostomya sp.) in 92.4% of trees, although placement of this subclade across the ingroup varies, as do the topologic positions of other freshwater species. Freshwater and marine corbulids also are morphologically distinct from each other, a factor that likely reduced the robustness of our phylogenetic results. By combining these results with paleoecologic, stratigraphic, and morphologic data, we infer that freshwater corbulids arose once within the family, prior to the Cenozoic, with three distinct freshwater lineages present at their first appearance in the late Paleocene of North America. Within the Miocene Pebas system of South America, we reconstruct supralimital morphologic evolution within three lineages as freshwater taxa became variously adapted to the fluid, dysoxic muds characterizing lake-bottom facies representative of the Pebas lacustrine system. In addition, corbulids apparently successfully coped with high predation pressures from co-occurring shell-crushing predators. Finally, we consider that freshwater Corbulidae were primarily fluvial taxa throughout their geologic history, with a relatively ephemeral radiation within the Pebasian lake system, thus making the Pebasian system a cradle of diversity for several corbulid lineages.

Knowing the structure and branching order of the evolutionary tree is essential to many studies of the evolutionary process and the history of life. Most attempts to reconstruct evolutionary trees are based on rectangular matrices, in which coded states are assigned for each character and each taxon under study. Everyone who has tried to infer the pattern of evolution from such character-taxon matrices is confronted with a great deal of “noise” and uncertainty. Often, there are multiple “most parsimonious trees” (those requiring the fewest evolutionary steps), which often differ significantly in topology; and many nodes in such trees are poorly supported; that is, there are contradictions and inconsistencies in the data. Some of this noise arises for reasons well known to phylogeneticists. Character states may arise multiple times, evolve in parallel, or reverse (homoplasy); taxa vary insufficiently among themselves; and diversification may be so rapid that data are insufficient to detect the order of branching. Polytomies are regarded as a nuisance rather than as real information. Characters may be coded inappropriately, so that there are too many redundancies, inapplicable character states, and multistate characters, as well as dependence among characters. Moreover, it is highly debatable whether evolution is parsimonious, and there are conflicts about how to establish the polarity of characters (primitive vs. derived). (For reviews and discussion see Sober 1988; Huelsenbeck and Crandall 1997.)

In the past fifteen years, the biological interpretation of vertebrate fossils has been markedly advanced by the application of experimental techniques in functional morphology, biomechanical and allometric modelling, more rigorously-derived empirical correlations between form and function in living species, and studies of modern community structure and of the taphonomic processes by which fossil assemblages are formed. Traditional paleontological methods of phylogenetic analysis are under attack by the advocates of cladistics; the resulting debate is leading to increased rigor in paleontological systematics. Paleobiogeographical studies of vertebrates has become an extremely active area of research due to the need to reanalyze past distribution patterns in the light of plate tectonics.

New information has extended the fossil record of vertebrates back to the Late Cambrian, but little more is known about the nature of the earliest vertebrates or of the pre-vertebrate ancestors. Much has been learned of the fishes antecedent to tetrapods but little of the earliest tetrapods. The early reptiles are now well known, but the nature of their amphibian ancestors is uncertain. Restudy of Archaeopteryx has suggested a dinosaurian ancestry for birds. The origin and early evolution of mammals is the subject of extensive research as a result of greatly augmented recent collecting. The most active and innovative research is on the biology of dinosaurs, especially on their physiology. Among Cenozoic mammals, the fossil record of primates has been greatly expanded and knowledge of primate history, from the basal prosimians of the Paleocene to early man, is increasing rapidly.

Research into the relationship between leaf form and climate over the last century has revealed that, in many species, the sizes and shapes of leaf characters exhibit highly structured and predictable patterns of variation in response to the local climate. Several procedures have been developed that quantify covariation between the relative abundance of plant character states and the states of climate variables as a means of estimating paleoclimate parameters. One of the most widely used of these is the Climate Leaf Analysis Multivariate Program (CLAMP). The consistency, accuracy and reliability with which leaf characters can be identified and assigned to CLAMP character-state categories is critical to the accuracy of all CLAMP analyses. Here we report results of a series of performance tests for an image-based, fully automated at the point of use, leaf character scoring system that can be used to generate CLAMP leaf character state data for: leaf bases (acute, cordate and round), leaf apices (acute, attenuate), leaf shapes (ovate, elliptical and obovate), leaf lobing (unlobed, lobed), and leaf aspect ratios (length/width). This image-based system returned jackknifed identification accuracy ratios of between 87% and 100%. These results demonstrate that automated image-based identification systems have the potential to improve paleoenvironmental inferences via the provision of accurate, consistent and rapid CLAMP leaf-character identifications. More generally, our results provide strong support for the feasibility of using fully automated, image-based morphometric procedures to address the general problem of morphological character-state identification.

Using the diameters of the channels at branch points, we quantitatively test three alternative models of the function of astrorhizae in stromatoporoids. The distribution of diameters at branch points is significantly different from the distribution that would be predicted from models of either a diffusive function or a bulk-flow system in which resistance to flow was constant at all levels of the branching hierarchy. The distribution of channel diameters is virtually identical to that predicted by a model (Murray's law) that simultaneously minimizes resistance to flow and some volume-related cost function. Astrorhizae thus carried a bulk flow of fluid and can be inferred to have been lined with cellular elements; the exchange sites associated with the fluid-transport system were distributed throughout the soft tissues of the stromatoporoid animal. The most parsimonious hypothesis of function, that the fluid-transport system was associated with suspension feeding, implies strong similarities between the structure of the stromatoporoid animal and living sponges.

Studies of the fates of clades during mass extinctions at the end of the Cretaceous and near the Cambrian-Ordovician boundary have indicated that geographically widespread groups have a much higher probability of survival than narrowly distributed groups. If biogeographic distribution does play a significant role in influencing the outcome of extinction events, then geographic variability in extinction intensity should be mirrored by patterns of endemism. A comparison of data for the terminal Cambrian extinction indicates that survival of trilobite families in Kazakhstan was significantly greater than in North America. As predicted, the Kazakh sequence was characterized by a significantly larger number of pandemic families. The Baltic Province, which includes Scandinavia, England, and Wales, was composed entirely of pandemic groups and did not suffer any trilobite extinction at the family level. The results are consistent with the suggestion that latitudinal variation in extinction magnitude may be the result of differences in biogeographic structure.

Crura, the calcareous support structures of the lophophore in rhynchonellide brachiopods, have historically been used to justify higher-level rhynchonellide classification and reveal major evolutionary lineages within rhynchonellides. Seventeen crural types have been described and categorized into four groups based on variation in overall structure and cross-sectional shape, but not evaluated in a quantitative or comprehensive manner. Heterochrony has been hypothesized to play a role in the evolutionary transitions among some types, but the structural, developmental, and phylogenetic context for testing these hypotheses has not yet been established. In this study, we use three-dimensional geometric morphometric techniques to quantify morphological disparity among all six crural morphs in Recent adult rhynchonellides, with the goal of delineating more objective criteria for identifying and comparing crural morphs, ultimately to test hypotheses explaining morphological transformations in ontogeny and phylogeny. We imaged the crura of seven Recent rhynchonellide species, using X-ray computed microtomography. We used landmarks and semi-landmarks to define the dimensions and curvature of the crura and the surrounding hinge area. Procrustes-standardized landmark coordinates were analyzed using a principal component analysis to test the discreteness of the individual crural morphs and named groups of morphs, and to identify features that vary most among the crural configurations.

Our results demonstrate that microCT imaging techniques provide novel ways to investigate the morphology of small features that may be otherwise impossible to quantify using more conventional imaging techniques. Although we predicted overlap among crural morphs in the 3-D shape space, the principal component analyses suggest that five of the six crural morphs differ distinctly from one another. Some but not all previously designated crural groups appear to exhibit morphological cohesion. This study establishes a quantitative morphological foundation necessary to begin an investigation of the phylogenetic significance of ontogenetic changes in crura, which will allow hypotheses of heterochrony to be tested.

Growth rhythms are described in the accretionary skeletons of Rafinesquina alternata, a Late Ordovician brachiopod from southeastern Indiana. Contiguous growth increments widen and narrow in repeating series, giving the appearance of adjacent clusters of increments. Fourier analyses of growth increment widths and counts of the number of increments within individual clusters yield similar periodicities. Increments vary in width over a period of approximately 19 increments, modulated with a lower amplitude oscillation of 27 increments. The number of increments per cluster falls into two groups; clusters having between 8 and 17 increments outnumber those having between 18 and 30 increments.

All specimens were obtained from a Maysvillian facies of the Dillsboro Formation, previously inferred to represent a shallow subtidal environmental setting. The growth periodicities described here are consistent with this interpretation. The intensity of tidal parameters such as emersion-immersion cycles, substrate shifts, changes in nutrient supply or in oxygen tension declines with depth as would the number of growth increments added each month in response to these factors. Thus, for these specimens, the maximum number of increments per cluster probably approximates the true number of “tidal” days in the Late Ordovician synodic month (period between full moons).

The paleoecological model derived from these analyses can be used in future studies to predict the rate of the earth's rotation and the motion of the moon in the Late Ordovician and, equally importantly, to evaluate the limits of uncertainty of such studies.

Among extant crinoids, the feather stars are the most diverse and occupy the greatest bathymetric range, being especially common in reef environments. Feather stars possess a variety of morphological, behavioral and physiological traits that have been hypothesized to be critical to their success, especially in their ability to cope with predation. However, knowledge of their predators is exceptionally scant, consisting primarily of circumstantial evidence of attacks by fishes. In this study the question whether regular echinoids, recently shown to consume stalked crinoids, also consume feather stars is explored. Aquarium observations indicate that regular echinoids find feather stars palatable, including feather stars known to be distasteful to fish, and that regular echinoids can capture and eat live feather stars, including those known to swim. Gut-content analyses of the echinoid Araeosoma fenestratum (Thomson, 1872), which is commonly observed with large populations of the feather star Koehlermetra porrecta (Carpenter, 1888) in video transects from marine canyons off the coast of France, revealed elements of feather stars in the guts of 6 of 13 individuals. The high proportion of crinoid material (up to 90%), and the presence of articulated crinoid skeletal elements in the gut of A. fenestratum, suggest that these echinoids consumed at least some live crinoids, although they may have also ingested some postmortem remains found in the sediment. Additionally, photographic evidence from the northeast Atlantic suggests that another regular echinoid, Cidaris cidaris (Linnaeus, 1758), preys on feather stars. Thus in spite of the broad suite of antipredatory adaptations, feather stars are today subject to predation by regular echinoids and may have been since the Mesozoic, when this group of crinoids first appeared.

Numerical simulations of neutral metacommunities are used here to predict the effects of growth and shrinkage of metacommunities, as well as their separation and merging caused by continental collision and rifting and their secondary eustatic effects. Although growth and shrinkage of metacommunities predictably change diversity, separating and merging metacommunities have counterintuitive effects. Separating and merging metacommunities change diversity within the individual areas, especially so for smaller areas, but they cause no change in total diversity of the system, contrary to previous predictions. The response times of metacommunities are likely to be geologically undetectable except for enormously large systems. These models can be used to predict the plate-tectonic effects on the diversity of terrestrial, coastal-marine, deep-marine, and oceanic-island systems. Of these, global and regional coastal-marine systems are the most acutely sensitive to the changes in area and fragmentation caused by plate tectonics. Oceanic-island systems also experience global and regional changes in diversity during supercontinent breakup and assembly, with the global effects driven by the changing length of volcanic arcs, and the regional effects also driven by secondary eustatic changes in shallow-marine area. Although individual terrestrial provinces or continents may experience substantial changes in diversity from rifting and collision, global terrestrial diversity should be unchanged except for the relatively modest contributions caused by the secondary eustatic effects on land area. These changes in diversity may be reinforced or counteracted by the changing latitudinal position of metacommunities.

Mean adult body mass of mammal taxa is a fundamental ecological variable. Variability in the distributions of body masses of a mammal fauna suggest variability in habitat structure. Mammal remains from the Marmes archaeological site in southeastern Washington State date between 13,200 and 10,400 b.p., during the Pleistocene–Holocene transition (PHT). Known environmental history prompts the expectations that the Marmes PHT mammal remains should represent greater species richness and a larger array of body-mass sizes than modern faunas in the Marmes locale and in open shrub-steppe habitats, and lower species richness and a smaller array of body-mass sizes than modern faunas in closed forest habitats; species richness and the array of body-mass sizes should be similar to that for a mixed habitat of cool shrub-steppe with scattered conifers. The Marmes PHT cenogram meets these expectations. Body-mass clumps displayed by the Marmes PHT mammal fauna fall between those of closed forests and open shrub-steppe habitats in terms of clump richness and breadth, and in terms of gap width. Marmes PHT body-mass clumps are very similar to those for the mixed habitat. Cenograms and body-mass clumps confirm conclusions drawn 40 years ago that the Marmes PHT habitat was much like that of today but cooler and with more plant biomass and greater structural diversity than today.

Faunal similarity among regions is often used as a means of identifying regions of endemism in fossil faunas. At least two large-scale taphonomic effects can affect apparent faunal similarity: stratigraphic and facies mismatching. In stratigraphic mismatching, an unconformity represents removal of most or all of a complete assemblage zone in one region, and the constituent taxa are mistakenly interpreted as having never inhabited that region. In facies mismatching, environmental differences between two regions (possibly unrecognized) cause the inference of a barrier that never existed. The two types of mismatching can work in concert if a facies was originally represented in a single stratigraphic interval that has been completely removed from one region. Analysis of faunal similarity via multivariate analysis of individual localities, coupled with comparison of the regions as single samples, may indicate mismatching if the results differ significantly.

We view these two problems as part of a suite of taphonomic effects that are not evident in paleobiological analyses of smaller geographic scope. First, there is ambiguity in the notion of “barrier,” even when a candidate is obvious. Second, barriers in paleobiogeography are often hidden and must be inferred from their effects rather than observed. Third, stratigraphic and facies mismatching produce effects on regional faunas similar to those produced by barriers. Anyone using barriers to explain faunal disruptions should address these three points.

Upper Eocene faunas of central Florida seem taxonomically distinct from those of the remainder of the Gulf Coastal Plain. This has historically been attributed to a known paleogeographic feature, the Suwannee Strait, which acts as a barrier. The amount of dissimilarity of the echinoid faunas is greater than the amount predicted as a result of sampling problems. Comparison of the results of multivariate and whole-region analyses suggests that mismatching of the two faunas, rather than a true barrier, causes the distinction. Principally facies, but also strata are mismatched. Early Late Eocene faunas inhabited terrigenous sands to the north of the strait and carbonate sands to the south and show the highest distinctiveness. Middle Late Eocene faunas inhabited primarily carbonate sands to the south and both carbonate sands and muds to the north. Overall similarity is higher for both local and regional analyses, and the faunas of northern sands are more similar to those of the southern region than to the northern mud faunas. Overall similarity across the strait is highest in the late Late Eocene strata when both regions had carbonate mud and sand facies. The faunas exhibit greater similarity within facies than they do within regions. Upper Late Eocene strata are poorly preserved north of the strait because of post-Eocene erosion.

Oceanic anoxic events (OAEs) are contemporaneous with 11 of the 18 largest Phanerozoic extinction events, but the magnitude and selectivity of their paleoecological impact remains disputed. OAEs are associated with abrupt, rapid warming and increased CO2 flux to the atmosphere; thus, insights from this study may clarify the impact of current anthropogenic climate change on the biosphere. We investigated the influence of the Late Cretaceous Bonarelli event (OAE2; Cenomanian/Turonian stage boundary; ~94 Ma) on generic- and species-level molluscan diversity, extinction rates, and ecological turnover. Cenomanian/Turonian results were compared with changes across all Cretaceous stage boundaries, some of which are coincident with less severe OAEs. We found increased generic turnover, but not species-level turnover, associated with several Cretaceous OAEs. The absence of a species-level pattern may reflect species occurrence data that are too temporally coarse to robustly detect patterns. Five hypotheses of ecological selectivity relating anoxia to survivorship were tested across stage boundaries with respect to faunality, mobility, and diet using generalized linear models. Interestingly, benthic taxa were consistently selected against throughout the Cretaceous regardless of the presence or absence of OAEs. These results suggest that: (1) the Cenomanian/Turonian boundary (OAE2) was associated with a decline in molluscan diversity and increase in extinction rate that were significantly more severe than Cretaceous background levels; and (2) no differential ecological selectivity was associated with OAE-related diversity declines among the variables tested here.