The EDTA-soluble fractions extracted from rostra of two late Cretaceous belemnites (Cephalopoda), Gonioteuthis and Belemnitella, were divided into an humic acid and a fulvic acid fraction (the latter was absent in Belemnitella). The extracts are compared with preparations from shells of two recent cephalopods, Nautilus pompilius and Sepia officinalis. Use was made of immunology, amino acid analysis, pyrolysis mass spectrometry and some other techniques.

The fulvic acid fraction of Gonioteuthis, a mixed peptide-like saccharide-like substance, produced confluent immunodiffusion patterns with an EDTA-soluble Nautilus extract against anti-Nautilus rabbit serum. The humic acid of Gonioteuthis did not contain D-alloisoleucine and its amino acid composition was very similar to that of the EDTA-insoluble fraction of Nautilus. This humic acid was enriched in polyphenol, which may be due to chemical reaction of peptides and carbohydrates during diagenesis. It is concluded that both fractions of Gonioteuthis are original belemnite materials that have undergone only minor alterations during diagenesis.

This is an exploratory study of biochemical compounds derived from fossils, with particular emphasis on immunological methods.

The Cambrian Radiation marks the appearance of representatives of virtually all major skeletonized phyla in the fossil record and clearly represents a fundamental episode in the history of life. Furthermore, the tempo and mode of this evolutionary event have been the subject of intense debate. One area that has been debated is how so many phylum-level body plans can have evolved in such a geologically brief period. Some have argued that there was enhanced morphological flexibility and fewer evolutionary constraints at this time, leading to greater morphological disparity of Early Cambrian faunas. Others have claimed that this is not true because the evolution of most of the animal phyla significantly predates the radiation or because they failed to detect a signature of decreasing morphological disparity through time. At present, the higher-level patterns of diversification during this time period and the relevant implications for Early Cambrian uniqueness are areas of active research interest and debate. Recognizing this debate, we used both a phylogenetic and a morphometric framework to study whether there is a signature of increasing morphological constraint and decreasing flexibility through time within one of the clades that is a significant constituent of the Early Cambrian biota, specifically, the olenelloid trilobites. In this species-rich clade, we found no evidence that morphological changes were becoming either increasingly constrained or less flexible in one of the dominant Early Cambrian metazoan clades as it passed through the Cambrian Radiation.

Fundamental to the interpretation of bone-bearing faunal deposits is an understanding of the taphonomic processes that have modified the once living fossil community. An often neglected source of bias is that of climate-averaging, which occurs when the duration of bone accumulation exceeds the duration of an individual climatic episode. Tropical and subtropical climate change is dominated by precessional cyclicity (~21,000 year cycle), which controls monsoon rainfall intensity and thus plant communities over time. Under a climate-averaging scenario, the paleoecological characteristics of a faunal deposit represent an amalgamation of more than one phase of the precessional cycle. We investigate the degree of climate-averaging in Plio-Pleistocene bone breccias from South Africa by comparing stable isotope measurements of fossil enamel with the evidence from high-resolution speleothem paleoclimate proxies. We conclude that each of the four faunal assemblages studied are climate-averaged, having formed over a time period in excess of one-third of a precessional cycle (~7000 years). This has implications for the reconstruction of hominin paleoenvironments and estimates of Plio-Pleistocene biodiversity. We hypothesize that climate-averaging may be a common feature of tropical terrestrial vertebrate assemblages throughout the Cenozoic and Mesozoic.

Chitin occurs in a variety of invertebrates, especially in arthropod cuticles, but is rarely reported in the fossil record. Although it has been detected in fossils as old as Middle Cambrian and Silurian, the majority of records come from much younger, Cenozoic deposits. In this paper, we report the preservation of chitin in Early Jurassic neritimorph gastropod egg capsules deposited in bivalve shells from prodelta-deltafront and nearshore paleoenvironments of the Holy Cross Mountains, Poland. We used a number of analytical methods to confirm the presence of chitin preserved in these ancient fossils. This is the first record of chitin preservation in Mesozoic deposits that, interestingly, do not follow the conventional Konservat-Lagerstätten manner of preserving soft-bodied and non-biomineralized organisms. We believe that deltaic settings characterized by episodic, high input of fluvial deposits, oligohaline conditions, and oxygen-poor microenvironment within the sediment—as well as early cementation of sediment infilling the shells—were crucial for chitin preservation. The preservation of chitin in such recalcitrant structures as egg capsules and deposits that formed outside conventional Konservat-Lagerstätten conditions renders it likely similar deposits may yield promise for discoveries of similar biological macromolecules.

Documenting past environmental disturbances will provide a very incomplete explanation of extinctions until more data on intrinsic (e.g., phylogenetic) responses to disturbances are collected. Taxonomic selectivity can be used to infer phylogenetic inheritance of extinction-biasing traits. Selectivity patterns among higher taxa, such as between mammals and bivalves, are well documented. Selectivity patterns among lower taxa (genus, species) have great potential for understanding the dynamics underlying higher taxic turnover. Two echinoid data sets, of fossil and living taxa, indicate that species extinctions do not occur randomly within genera. Reverse rarefaction estimates of past species extinction rates assume random species extinction within higher taxa, so these widely cited extinction estimates may be inaccurate. Revised estimates based on a simulated curve imply that past species extinctions rates may be 6%–15% lower than previously cited. Possible causes for the observed selectivity patterns are discussed. These include nonrandom phylogenetic nesting of species with traits often cited as enhancing extinction vulnerability, into certain taxa. Such traits include low abundance, large body size, narrow niche breadth, and many others. Phylogenetic nesting of extinction-biasing traits at many taxonomic levels does not predict that a dichotomy of mass-background selectivity based on a few traits will occur. Instead, it predicts patterns of selectivity at many taxonomic levels, and at many spatio-temporal scales.

It has been claimed that articulate brachiopods are excluded from turbulent environments by the weakness of their pedicle attachment. The evidence does not support this conclusion. Experiments on the force required to remove brachiopods from their substrate show that removal force often equals that of bivalve mussels from turbulent environments having the same valve area. Intertidal brachiopods probably occupy cryptic habitats to avoid desiccation, not to avoid waves. Subtidal brachiopods are abundant in tidal currents of 4 knots (2 m/sec) where they are usually attached in exposed positions.

Although paleoecologists often assume that the size of the pedicle foramen is directly proportional to attachment strength, the relationship is not precise. Experimental detachment of brachiopods resulted from failure of the substrate, pedicle-substrate attachment, pedicle, or pedicle adjustor muscles, in approximately equal frequencies. At the same attachment strength, the foramen in Terebratalia is four times the size of the foramen in Hemithiris.

Although subapical foramina are supposedly unable to enlarge as the brachiopod grows, the subapical foramen of Hemithiris appears to do so.

Small terrestrial vertebrates are not capable of maintaining a constant body temperature (±2°C) without a relatively high metabolism. The amount of temperature variability during bone growth can be determined using oxygen isotopes from bone phosphate because fractionation of oxygen isotopes between body fluid and bone phosphate is dependent upon temperature. Fluctuation of body temperature during the early phase of growth in juvenile ectotherms should result in high intra- and interbone isotopic variability, whereas juvenile endotherms should have low isotopic variability resulting from the maintenance of homeothermy. Analyses of juvenile Hypacrosaurus individuals indicate a pattern of low isotopic heterogeneity suggestive of endothermy.

Molecular data are becoming an indispensable tool for the reconstruction of phylogenies. Fossil molecular data remain scarce, but have the potential to resolve patterns of deep branching and provide empirical tests of tree reconstruction techniques. A total evidence approach, combining and comparing complementary morphological, molecular and stratigraphical data from both recent and fossil taxa, is advocated as the most promising way forward because there are several well-established problems that can afflict the analysis of molecular sequence data sometimes resulting in spurious tree topologies. The integration of evidence allows us to: (1) choose suitable taxa for molecular phylogenetic analysis for the question at hand; (2) discriminate between conflicting hypotheses of taxonomic relationship and phylogeny; (3) evaluate procedures and assumptions underlying methods of building trees; and (4) estimate rates of molecular evolution in the geological past. Paleontology offers a set of independent data for comparison and corroboration of analyses and provides the only direct means of calibrating molecular trees, thus giving insight into rates of molecular evolution in the geological past.

Numerous environmental and intrinsic biotic factors have been sought to explain patterns in diversity and turnover. Using taxonomically vetted and sampling-standardized data sets of more than 50,000 taxonomic occurrences in the Paleobiology Database (PaleoDB) we tested whether habitat breadth predicts genus durations and diversity dynamics of marine Mesozoic bivalves, and whether this effect is independent of the well-known positive relationship between geographic range and longevity. We defined the habitat breadth of a genus as a function of its realized ranges in water depth, substrate type, and grain size of the substrate. Our analysis showed that mean values of extinction and origination rates are significantly higher for narrowly adapted genera compared to broadly adapted genera, with differences being evident in all analyzed stratigraphic intervals. Linear models showed that both geographic range and habitat breadth have an independent effect on genus durations and on diversity dynamics. These results reaffirm the role of geographic range and furthermore suggest that habitat breadth is an equally important key predictor of extinction risk and origination probability in Mesozoic marine bivalves. Habitat generalists, regardless of their geographic range, are generally less prone to extinction. Conversely, widely distributed genera that are more specialized may be more endangered than one would expect from their geographic range alone. Extinction rates tend to be higher for specialized genera in both background and mass extinctions, suggesting that wide habitat breadth universally buffers against extinction. The trajectories of origination rates through time differ from those of extinction rates. Whereas there is no pronounced ecological selectivity in origination in the Triassic and most of the Jurassic, Cretaceous origination rates are higher for specialized genera. This may best be explained by diversity-dependence. When diversity levels reach a critical point a further increase in diversity is achieved by elevated origination rates of more specialized forms.

Faunal abundance and fossil size and shape data from microvertebrate localities in the Upper Cretaceous Judith River Formation of south-central Montana illustrate that even when located in similar sedimentary facies, concentrations of vertebrate microfossils may exhibit strikingly different taphonomic profiles. Degrees of microfossil size and shape sorting may vary even among sites from the same sedimentary facies. In some instances, such variations may make it impossible to disregard taphonomic causes for differences sites exhibit in paleofaunal abundances. To limit the possibility that taphonomically generated faunal differences might be mistaken for true differences in paleoecology, comparisons of paleofaunal abundances should be restricted to microvertebrate sites both from similar sedimentary facies and with similar profiles of fossil sizes and shapes.

The longevity of genera living at the edge of the early Ordovician North American plate is related to their position on the former shelf, that is preserved in richly fossiliferous sections of Spitsbergen. Genera which endure for a long period (SO Myr or more) tend to be confined to shallow-water sites associated with algal mounds at the edge of the carbonate platform (illaenid-cheirurid community type) or deep-water sites with low oxygen concentration, and probably beneath the thermocline (olenid community type). The latter includes “conservative” forms that have survived from the Cambrian. The long ranges of the olenid community type are attributed to the adaptation of only a few genera to a highly specific, stressed environment. The endurance of the illaenid-cheirurid genera is due to the persistence of the tropical shelf-edge habitat, with high predictability of resources, to which the Ordovician forms became adapted early on. The nileid community type, which occurred between the platform edge and olenid-bearing environment, was the site of rapid generic turnover and may have been the source for subsequent recruitment into other environments. The inner shelf also had rapid generic replacement, dominated by a single family, the Bathyuridae. Rapidity of evolution in this site may be due to the spatial heterogeneity of the epeiric environment, as in the Devonian phacopids studied by Eldredge (1974).

During the latest Cretaceous and the Paleocene in western North America, disappearance rates for mammalian genera track appearance rates, both reaching their peak in the early Paleocene (Puercan) following the extinction of non-avian dinosaurs. Some of the disappearances during this time were pseudoextinctions that resulted when ancestral species disappeared during speciation.

Species-level cladistic analyses and a well-constrained biostratigraphic framework are required to study this form of pseudoextinction. Cladistic analyses show that monophyly cannot be established or rejected for some species because these species lack autapomorphies (uniquely derived character states) that unite their constituent members. Such taxa, termed metaspecies, are potential ancestors to species and higher clades with which they share a node in the cladogram.

A hypothetical species-level cladistic analysis coupled with three different hypothetical biostratigraphies shows how different models of speciation (bifurcation, budding, or anagenesis) result in very different patterns of true versus pseudoextinction. Depending on the speciation model, true extinction can be overestimated by as much as a factor of four, raising the specter of mass extinction. Species-level studies for three early Tertiary mammalian taxa—taeniodont eutherians, taeniolabidid multituberculates, and periptychid ungulates—use the same procedures. They show that almost 25% of disappearances during the early Paleocene (Puercan) for species in the analysis were pseudoextinctions of metaspecies. Budding and anagenetic-like peripatric speciation, but not bifurcation, are seen in the three examples.

Equating disappearance to true extinction can profoundly affect interpretations of faunal turnover, especially during mass extinctions or major faunal reorganizations. Some authors use pseudoextinction to describe the taxonomic rather than evolutionary disappearance of nonmonophyletic groups. Pseudoextinction, as used here refers only to the evolutionary disappearance of metaspecies via speciation. Both usages seem appropriate but should not be confounded.

The potential of the ichnofossil record for exploring the evolution of behavior has never been fully realized. Some of this is due to the nature of the trace fossil record itself. Equally responsible is the separation of ichnology from the relevant areas of modern behavioral biology. The two disciplines have virtually no concepts, methods, or literature in common. The study of animal behavior and its evolution is thus bereft of the rich data and insights of ichnologists.

One potential pathway forward is for ichnologists to adopt and adapt the movement ecology paradigm proposed several years ago by Ran Nathan and colleagues. This approach views movement as resulting from interactions of the organism's internal state, its movement abilities, and its sensory capabilities with each other and with the external environment. These interactions produce a movement path. The adoption of this paradigm would place trace fossil studies in a far wider common context for the study of movement, while providing the dimension of the evolution of movement behavior in deep time to neontological studies.

A second component of this integration would be for paleontologists to develop a taphonomy of behavior that places in a phylogenetic context the range of possible behaviors that organisms can carry out and assesses the potential of each of these behaviors in leaving a diagnostic trace. Parallel to other taphonomic concepts, this approach assesses the preservation potential of particular behaviors; behavioral fidelity is the extent to which trace fossils preserve these original behavioral signals.

Exploring patterns in the evolution of seed plant reproductive morphology within a functional context offers a framework in which to identify and evaluate factors that potentially drive reproductive evolution. Conifers are a particularly useful group for studies of this kind because they have a long geologic history and their reproductive organs are borne on separate structures with discrete functions. Multivariate analysis of morphological data collected from pollen-producing and seed-producing cones of Paleozoic, Mesozoic, and extant conifer species shows that seed cones underwent a significant expansion of morphological diversity that began during the Early-Middle Jurassic and has continued into the present day. In contrast, pollen cones show significantly lower levels of morphological diversity and exhibit similar basic morphologies throughout conifer evolutionary history. The increase in seed cone diversity through time is primarily the result of two novel structural and organizational features that evolved independently in different conifer families during the Mesozoic: robust, tightly packed cones in members of Araucariaceae, Cupressaceae sensu lato, and Pinaceae, and highly reduced, fleshy cones or solitary seeds in Podocarpaceae, Taxaceae, and some members of Cupressaceae sensu stricto. In extant conifers, these cone morphologies are associated with species that have strong interactions with vertebrate seed predators, seed dispersers, or a combination of both. This suggests that increases in the strength and complexity of biotic interactions in the Jurassic and Cretaceous were a primary driver of conifer reproductive evolution, and that patterns of character evolution relate to the increasing importance of cone tissue in seed protection and seed dispersal through time.

A variety of means, including forelimb proportions and shell bone histology have been used to infer the paleoecology of extinct turtles. However, the height-to-width ratio of the shell (as a one-parameter shell model) has been dismissed because of its unreliability, and more complex aspects of shell geometry have generally been overlooked. Here we use a more reliable, three-parameter geometric model of the shell outline in anterior view as a means to assess turtle paleoecology. The accuracy of predictions of extant turtle ecology based on our three-parameter shell model is comparable to that derived from forelimb proportions when distinguishing between three ecological classes (terrestrial, semiaquatic, and aquatic). Higher accuracy is obtained when distinguishing between two classes (terrestrial and non-terrestrial), because the contours of aquatic and semiaquatic turtles are often very similar. Our model classifies Proterochersis robusta, a stem turtle from the Late Triassic of Germany, as non-terrestrial, and likely semiaquatic. Our method, combined with inferences based on limb proportions, indicates a diverse range of ecotypes represented by Late Triassic stem turtles. This implies that the ecological diversification of stem-group turtles may have been rapid, or that a substantial period of currently cryptic diversification preceded the first fossil appearance of the turtle stem lineage during the Late Triassic.

The mineralized hard parts of Nautilus, including the mandibles, statoconia, and uroliths, are more complex with respect to ultrastructure, mineralogy, trace element, and isotopic composition than was previously recognized. X-ray diffraction and amino acid composition analyses of the organic structural hard parts (siphuncle tube, shell wall and septum organic matrix, mandibles, and muscle tendon sheath) show that two different chitin-protein complexes are utilized by Nautilus. The mandible mineral hard parts are particularly complex, with five different minerals present in various locations. A comparison of the statoconia of Nautilus species with the statoliths of dibranchian cephalopods reveals an evolutionary trend in which carbonate is substituted for phosphate. This study also shows that Nautilus uses a number of different crystal-forming processes for constructing its hard parts. The data presented here provide a broad spectrum of information, which, when applied to the fossil counterparts, can be utilized for improving our understanding of ancient nautiloid biology.

Implosion of 47 epoxy-sealed shells of Nautilus pompilius indicates that very young shells (20-30 mm diameter at last septum) can withstand 136 atm pressure (= 1360 m equivalent depth), but intermediate and mature shells (30-145 mm diameter) implode at approximately 30–70 atm (= 300–700 m). Strain gages attached to the shells during implosion tests show that the flank and venter are compressed by external pressure, but the septum undergoes tension and structurally should be the most vulnerable to hydrostatic pressure. Overall shell strength is substantially less than reported values of tensile strength (approximately 364 atm) and compressive strength (approximately 1561 atm) of the constituent shell material, nacre. Shell strength as a depth limiting factor appears to be controlled by shell and septal architecture and by stress concentrators such as flaws and microstructural inhomogeneity. The wide range in implosion values and lack of strong correlation between such parameters as septal thickness and implosion pressure indicate that determination of depth ranges for fossil cephalopods may be difficult.

Morphological analysis of four higher taxa of fossil marine invertebrates shows that, over the history of paleontology, there is no general tendency for morphologically extreme or modal species and genera to be described preferentially early or late. Reconstructing the expected evolutionary sequences of morphological disparity that would have been estimated at various times during the past century and a half reveals features that are sensitive to sampling (for example, peak trilobite disparity in the Ordovician, peak of post-Paleozoic crinoid disparity in the Triassic, and peak blastoid disparity in the Permian), as well as more robust features (for example, increase in trilobite disparity from the Cambrian to the Ordovician, continued increase in trilobite disparity despite a drop in taxonomic diversity after the Early Ordovician, decrease in blastoid disparity from the Devonian to the Carboniferous, and increase in crinoid disparity from the Jurassic to the Cretaceous followed by decline during the Cretaceous). Although we still have much to learn about the evolution of form, in many respects our view of the history of biological diversity is mature.

Quantitative systems for describing naticid borehole position in bivalve and gastropod prey are presented. Pleistocene nassariid prey were bored at 53% of the distance up the shell and 8° to the right of the columella. A similar stereotyped position was found for Recent prey. The longitudinal shell axis and the aperture of the gastropod shell are proposed as primary cues in orienting prey for boring.