In cheilostome bryozoans with rigidly arborescent growth habits, resistance to breakage under forces generated by flowing water or collision with moving objects depends on the strength and design of supporting skeletal parts. To investigate the abilities of modern and Tertiary cheilostomes to resist breakage, we measured bending strength in fresh, preserved, and fossil material; measured drag on actual and model colonies; and calculated stress within colonies and breaking values under concentrated and uniform loads. Among nine modern, five Oligocene, and four Paleocene species, inferred live bending strength (24–85 MNm−2) and stiffness (42–65 GNm−2) appear to be species-specific properties overlapping those of similar mineral-organic composite skeletons of Mollusca. Unlike those of Mollusca, cheilostome skeletons appear isotropic in bending, with strength not clearly related to microstructure or composition. Bending strength and morphologic disposition of skeletal material combine to produce a wide range of abilities to resist breaking. At early growth stages, almost all species appear highly resistant; at later ones, some break at velocities <0.2 m sec−1 or concentrated loads <2 g, whereas others remain unbroken at velocities >2 m sec−1 or concentrated loads >100 g. A negative correlation between strength and resistance indicates that design is the more important factor in the abilities of these bryozoans to withstand forces of these kinds. An apparent trend toward more resistant designs from Paleocene to modern species is related chiefly to increasing rates at which branches thicken toward the colony base. The consequent change in design may even have permitted modern species to use weaker skeletal material while increasing their overall resistance.

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.

The known Cretaceous formicoids are better interpreted from morphological evidence as forming a single subfamily, the Sphecomyrminae, and even a single genus, Sphecomyrma, rather than multiple families and genera. The females appear to have been differentiated as queen and worker castes belonging to the same colonial species instead of winged and wingless solitary females belonging to different species. The former conclusion is supported by the fact that the abdomens of workers of modern ant species and extinct Miocene ant species are smaller relative to the rest of the body than is the case for modern wingless solitary wasps. The wingless Cretaceous formicoids conform to the proportions of ant workers rather than to those of wasps (Figs. 1–2) and hence are reasonably interpreted to have lived in colonies.

The Cretaceous formicoids are nevertheless anatomically primitive with reference to modern ants and share some key traits with nonsocial aculeate wasps. They were distributed widely over Laurasia and appear to have been much less abundant than modern ants.

Understanding the extent to which the reported fossil record reflects biological history, rather than preservational artifacts or other biasing factors, remains one of the central issues in the interpretation of the history of life on Earth. The development of large interactive paleontological databases, such as the Paleobiology Database (PBDB), allows detailed analyses of the patterns of occurrence, both regionally and globally, of taxa in the fossil record and makes possible testing hypotheses of the controls of the patterns. An analysis of data from the PBDB shows that most genera in the fossil record are rare, whereas a relatively small percentage of taxa account for a disproportionate share of the total occurrences. These ubiquitous taxa tend to be speciose and have long stratigraphic ranges. These patterns of occurrence might represent a true biological signal; it is also possible that they reflect taphonomic processes or are the result of taxonomic practice. In particular, common taxa may be taxonomic wastebaskets, i.e., residual and polyphyletic groups resulting from inadequate systematic attention and/or from taphonomic biases resulting in inadequate specimens being preferentially placed in particular genera. A conceptual model for the development of taxonomic wastebaskets suggests that these taxa should be speciose, widely distributed, common, and old (in terms of year of first description), and that they should be the nominate forms for higher taxa. Our analyses suggest that many of the common taxa in the PBDB are consistent with two or more of these expectations and are thus good candidates for being wastebaskets. These taxa are, however, only a small percentage of total genera. A more detailed examination of one group, early gastropods, indicates that possible wastebaskets still are present in a group that has received much recent systematic work. Given that likely wastebasket taxa are a small fraction of all genera, they probably have little effect on overall temporal patterns of generic richness. Their impact on other types of metrics, such as turnover rates or metrics of community diversity or biogeographic similarity, however, might be quite important.

The end-Permian mass extinction, the largest extinction of the Phanerozoic, led to a severe reduction in both taxonomic richness and ecological complexity of marine communities, eventually culminating in a dramatic ecological restructuring of communities. During the Early Triassic recovery interval, disaster taxa proliferated and numerically dominated many marine benthic invertebrate assemblages. These disaster taxa include the bivalve genera Claraia, Unionites, Eumorphotis, and Promyalina, and the inarticulate brachiopod Lingularia. The exact nature and extent of their dominance remains uncertain. Here, a quantitative analysis of the dominance of these taxa within the fossil communities of Panthalassa and Tethys benthic realms is undertaken for the stages of the Early Triassic to examine temporal and regional changes in disaster-taxon dominance as recovery progresses. Community dominance and disaster-taxon abundance is markedly different between Panthalassic and Tethyan communities. In Panthalassa, community evenness is low in the Induan stage but increases significantly in the Smithian and Spathian. This is coincident with a significant decrease in the relative abundance and occurrence frequency of the disaster taxa, most notably of the low-oxygen-affinity taxa Claraia and Lingularia. While the disaster taxa are present in post-Induan assemblages, other taxa, including two articulate brachiopod genera, outrank the disaster taxa in relative abundance. In the Tethys, assemblages are generally more even than contemporaneous Panthalassic assemblages. We observe an averaged trend toward more even communities with fewer disaster taxa in both Panthalassic and Tethyan assemblages over time.

The foraging paradigm of trace fossil theory has historically accorded random behavior to non-food-limited deposit-feeders and non-random behavior to food-limited feeders. A series of randomness measures derived from empirical modeling, simulation modeling, stochastic modeling and probability theory applied to foraging patterns observed in deep-sea bottom photographs from the Arctic and Antarctic yielded a behavioral continuum of increasing non-randomness. A linear regression of trace positions along the continuum to bathymetric data did not substantiate the optimal foraging efficiency-depth dependence model of trace fossil theory, except that all traces exhibited a greater optimization than that of simulated random foraging. It is hypothesized that optimization as evidenced by non-random foraging strategies represents maximization of the cost/benefit ratio of resource exploitation to risk of predation and that individual foraging patterns reflect an exploration response to the morphometry of a patchily distributed food resource. Differential predation and competition may account for the co-occurrence of random and non-random strategies within the same bathymetric zone.

In North America, hipparion horses consisted of four distinct genera, Hipparion, Neohipparion, Nannippus, and Cormohipparion, known from the medial Miocene to the late Pliocene, ca. 15.5–2 ma ago. The speciation mode is determined for 10 of the 16 valid hipparion species; 50% (5) seem to have resulted from anagenesis and 50% (5) from cladogenesis. Roughly contemporaneous species of the four hipparion genera seem to proceed through similar evolutionary stages of cranial and dental characters. The mean hipparion species longevity is 3.3 ma (N = 16); the mean generic longevity is 8.4 ma (N = 4). Only two species become extinct in less than 2 ma; extinction occurs at a rate of ca. 470 μmacarthurs for species with durations between 2 and 5 ma. For selected dental characters, mean evolutionary rates (for the 16 species) were between .08 darwins (d) and .03 d with crown height evolving most rapidly. Despite the commonly held notion that fossil horses evolved rapidly, these data suggest normal or average (horotelic) evolution for hipparions, which is not surprising because they were already established in the grazing adaptive zone during their history.

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.

A late Miocene population of Neohipparion cf. leptode from the Love Bone Bed of Florida was found to have 13 discrete annual age classes based on eruption sequences and tooth wear. The large sample size of individuals (minimum number = 229) allowed the estimation of standard population parameters, including age-specific mortality rates. The average age of death for a newborn was 3.5 years, but those individuals that survived the 64% juvenile mortality occurring during the first two years lived an average of 8.0 years. The potential longevity of this species of Neohipparion had not increased beyond that of known samples of Merychippus, despite increased crown height. The paleoenvironment of the collecting site was a wooded grassland savanna with annual wet and dry seasons. The Neohipparion population migrated into the area during the dry season but left in the wet season to give birth away from the collecting site.

Evidence that can be used to interpret the life habits of extinct organisms usually takes three forms: functional analysis, analogy with living organisms, and geological evidence. Independent quantitative tests for habit are rarely available. A theory of optimum eye design originally derived for living aquatic arthropods provides quantitative data that are used to test previous suppositions about the life habits of two Ordovician pelagic trilobites: that the telephinid trilobite Carolinites was epipelagic while the cyclopygid Pricyclopyge was mesopelagic. Optimum compound eye design theory uses measured lens diameters and interommatidial angles to determine the “eye parameter” (p), which can be used to gauge approximate optimum level of illumination for the eyes of these trilobites. The eye parameter provides an independent test for their relative paleobathymetry. Values of the eye parameter measured in the dorso-ventral direction across two eyes of Carolinites killaryensis utahensis were found to have medians of 2.13 and 3.24. Values measured in the antero-posterior direction have medians of 3.17 and 4.86 for the two eyes. Values measured in the dorso-ventral direction across two eyes of Pricyclopyge binodosa have medians of 4.23 and 4.98, while values measured in the antero-posterior direction have medians of 7.06 and 8.31. Eye parameters are higher in Pricyclopyge than in Carolinites, the difference statistically significant at p = 0.05. The eyes of Pricyclopyge are optimally designed for lower levels of illumination than are those of Carolinites. This accords with the previous interpretations of the former trilobite as mesopelagic and the latter as epipelagic.

Experienced workers in a field are unlikely to change their assessment of basic philosophical issues. Einstein for example never accepted the indeterminacy required by the Heisenberg uncertainty principle, despite the urgings of Max Born and virtually all others. Similarly even earlier, some older workers had refused to accept the young Einstein's theories. Now if issues in physics are not open and shut, they are even less so in paleontology. Possibly it is useful to probe the underlying basic philosophical background to our discipline. The purpose of the present article is to indicate that the inductive, deterministic approaches which have been used almost exclusively in paleontology have stringent limitations. For some types of problems, deductive stochastic processes are more interesting. If we can make the distinction of ‘few’ versus ‘many,’ then it is quite easy to consider deterministic explanations for the individual cases (the movement of individual molecules) but to reserve stochastic explanations for the ensembles of events (the movement of molecules too numerous to count).

A rate of extinction must specify both the amount of taxonomic change and the duration of that change. A consistent means of temporal measurement must be used to establish the duration. Terms used to describe the extinction, like catastrophic and gradual, should be quantitatively defined in relation to this framework.

Resolving rates of extinction in the stratigraphic record is limited by (1) the precision of chronostratigraphic correlations between individual sections and (2) the temporal completeness of litho- and biostratigraphic records in those sections. In this study, temporal completeness is estimated in eight of the best-known fluvial and pelagic sections spanning the K-T boundary. Temporal standardization is provided by correlations to the geomagnetic polarity time scale. A review of the literature documents lithologic criteria that might be used to infer the presence of a hiatus of some length at the biostratigraphically recognized K-T boundary. Rather than rejecting the presence of such gaps, this study argues that a hiatus of some length does exist and tries to estimate both a maximum limit for its duration and the probability that it represents a given duration below that limit.

Pelagic sections are probably more complete than fluvial ones. The section at Caravaca, Spain, can be expected to be the most complete, probably preserving sediment during each 10,000-yr interval of Chron 29R. The section in the San Juan Basin, New Mexico, is expected to be the most complete fluvial section, probably preserving sediment during each 100,000-yr interval of 29R.

Results indicate that our best-known marine and terrestrial sections spanning the K-T boundary should not be expected to document biologically catastrophic rates of extinction. A sufficiently precise means of global correlation that is independent of biostratigraphy and causal hypotheses is still unavailable, and it is very unlikely that the sections examined are complete at the 100-yr or biologically catastrophic level of precision. So, although catastrophic amounts of extinction might have occurred during the K-T transition, it seems unlikely that we can distinguish episodes of extinction lasting 100 yr or less from episodes lasting as long as 100,000 yr. Consequently, acceptance of catastrophic hypotheses based on these stratigraphic records seems improbably optimistic at this time.

The earliest Paleocene record of calcareous nannoplankton presents a unique opportunity to understand the evolutionary recovery of life from mass extinction. Nannoplankton were devastated at the Cretaceous/Paleogene boundary and their subsequent recovery can be studied in great detail because of their abundance in sediments, continuous stratigraphic occurrence, and near global distribution. Here we determine when and where new species of nannoplankton originated and how they dispersed following the Cretaceous/Paleogene mass extinction. Initially, we focus our efforts on North Pacific and South Atlantic deep sea sites with orbital age control to compare the precise timing and dynamics of the recovery between the locations. We then broaden our investigation to six sites from different basins and a variety of environments to study global patterns of the initial recovery. Our results show that many taxa in key Paleogene lineages originated in the North Pacific Ocean and that assemblages comprised primarily of new Paleogene taxa were not observed at other sites for several hundred thousand years. Survivors that were adapted to eutrophic post extinction conditions rapidly expanded in Southern Hemisphere sites where they dominated assemblages for most of the initial recovery. We therefore hypothesize that groups of survivors formed regionally incumbent assemblages in the Southern Hemisphere that limited diversification and dispersal of new Paleogene taxa. The end of survivor dominance correlates to the recovery of the biologic pump and subsequent decrease in surface ocean nutrient concentration 300–400 Kyr after the boundary. Only after survivors were removed did new Paleogene nannoplankton assemblages become abundant globally. Our results indicate that competition from regionally incumbent survivors was as an important control on the K/Pg recovery of nannoplankton.

The mammal fauna of the Willwood Formation, central Bighorn Basin, Wyoming, is ideal for paleoecological analysis because it is extensive, well studied, and continuously distributed over sediments representing the first 3 Myr of the early Eocene. The geology of the Bighorn Basin is also well known, providing a precise temporal framework and climatic context for the Willwood mammals. Previous analysis identified three “biohorizons,” based on simple counts of the first and last appearances of species. This study uses species diversity and appearance rates calculated from more extensive collections to approximate the ecological dynamic of the ancient fauna and assess whether the biohorizons were significant turnover events related to recently described climatic variation. Diversity and appearance data collected for this project are extensively corrected for uneven sampling, which varies by two orders of magnitude. Observed, standardized appearance and diversity estimates are subsequently compared with predicted background frequencies to identify significant variation. Important coincident shifts in the biotic parameters demonstrate that ecological change was concentrated in two discrete intervals ≤300 Kyr each that correspond with two of the original biohorizons. The intervals coincide with the onset and reversal of an episode of climate cooling identified directly from Bighorn Basin floras and sediments. Ecological changes inferred from the diversity and turnover patterns at and following the two biohorizons suggest short- and long-term faunal response to shifts in mean annual temperature on the order of 5–8°C.

In Baculites, a straight shelled ammonite, the constructional limits on shell shape resulting from the limited strength of nacre in tension are circumvented by a system of vaults in the phragmocone. Vaults bridge between regions of the phragmocone supported by the complex ammonite septal suture, and maintain the shell wall in compression when hydrostatic load induces bending moments. To determine how these vaults interact in the phragmocone to resist hydrostatic loading, measurements were made on a suite of Baculites specimens. In Baculites there is a statistically significant inverse relationship between circumferential curvature (radius of curvature) of the vaulted shell surface and the size of vaults spanning between sutural elements supporting the phragmocone. The inverse relationship between radius of curvature and the sizes of spans in this system of vaults results in the generation of comparable reactive forces at the ends of the vault spans where adjacent vaults interact. The equivalence of these reactive forces prevents the lateral displacement of the vault ends. Consequently, compressive stresses from adjacent vaults are superimposed on, and reduce, the tensional stress component of bending. Limiting tensile stress is of utmost importance in a lightweight shell composed of a brittle material such as nacre, which is strong in compression but weak in tension.

Baculites is particularly appropriate for this study because its straight shell is curved only in the circumferential direction, thus simplifying the problem. However, sutural patterns in coiled ammonites appear to be similarly constrained to produce vaults in the phragmocone which vary inversely in curvature and span size.

A global database of middle–upper Permian foraminiferal genera has been compiled from the literature for 75 Guadalupian and 62 Lopingian localities, grouped into 32 and 19 operational geographical units respectively. Cluster analysis reveals that five distinct Guadalupian provinces were reduced to four in the Lopingian, following the disappearance of the Eastern Panthalassa Province. Extinction magnitudes across the Guadalupian/Lopingian (G/L) boundary reveal that, in the remaining provinces, there is a strong regional variation to the losses at low paleolatitudes. The Central and Western Tethys Province experienced a markedly lower extinction magnitude, at both provincial and global levels, than the Eastern and Northern Tethys Province. Panthalassa experienced a high extinction magnitude of endemics, but a global extinction magnitude similar to that recorded in Central and Western Tethys. This regional bias is seen in both the fusulinacean and non-fusulinacean foraminifera, although fusulinaceans suffered much higher magnitudes of extinction. The regional selectivity also persisted during the subsequent Lopingian radiations, with the Central and Western Tethys Province recording the greatest magnitudes. Thus, of 35 new genera recorded globally from the Lopingian, 27 of these are recorded in Central and Western Tethys, compared to five and 12 genera respectively in Panthalassa and in Eastern and Northern Tethys. The Emeishan large igneous province erupted within the Eastern and Northern Tethys Province and may have been a factor in the high extinction–low radiation regime of this region. Regression (and consequent shallow-marine habitat loss) also appears to have been a significant factor. A major, but brief, late Guadalupian regression is best seen in those areas that suffered the greatest extinction losses.

Immunologically reactive protein substances were extracted from bone samples of an American mastodon (Mammut americanum), 10,200 yr old by radiocarbon dating, and from muscle samples of three woolly mammoths (Mammuthus primigenius), 10,000, 40,000 and 53,000 yr old, respectively. The mastodon samples contained significant quantities of the amino acids hydroxylysine and hydroxyproline, both of which are usually found in collagens and not in albumins. Using these products and other comparable extracts, as well as sera and purified collagens from modern elephants and other living mammals, as test antigens, immunological comparisons were carried out with the following antisera: rabbit anti-mastodon bone; chicken anti-mammoth muscle; chicken anti-elephant muscle; rabbit anti-elephant albumin and rabbit anti-elephant collagen, as well as with rabbit antisera to purified albumins and collagens of other mammals. For the first time, mastodon bone was found to have elephant-like proteins, which elicited antibodies that reacted strongly with collagen and serum proteins of extant elephants. Mammoth muscle strongly reacted with anti-elephant collagen and anti-elephant albumin, but the concentrations of the recoverable mammoth collagen and albumin decreased with increasing chronological age of the mammoth specimens. Nevertheless, in the immunological comparisons, the mammoth was closer to Asian (Elephas maximus) and African (Loxodonta africana) elephants than to the mastodon; in turn, the mastodon was closer to these elephantid species than to mammals outside the order Proboscidea.

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.

Major shifts in ecological dominance are one of the most conspicuous but poorly understood features of the fossil record. Here we examine one of the most prominent such shifts, the Ordovician shift from trilobite to brachiopod dominance of benthic ecosystems. Using an integrated database of high-resolution paleoecological samples and body size data, we show that while the average local richness and relative abundance of trilobites declined significantly through the Ordovician, the estimated standing biomass of trilobites, and by implication the amount of energy that they used, remained relatively invariant. This is attributable to an increase in the average body size of trilobite species in our data set, and especially to the widespread occurrence of the exceptionally large Middle-Late Ordovician trilobite genus Isotelus. Brachiopods increase in both mean body size and relative abundance throughout the Ordovician, so that estimates of brachiopod biomass and energetic use increase substantially between the Early and Late Ordovician. Although the data set includes a range of depositional environments, similar trends are observed in both shallow subtidal and deep subtidal settings. These results suggest that diversification of the Paleozoic Fauna did not come at the energetic expense of the Cambrian Fauna. The declining relative abundance of trilobites may reflect a combination of numerical dilution and the necessary energetic trade-offs between body size and abundance.

The preservation of compounds of biological origin (nucleic acids, proteins, carbohydrates, lipids, and resistant biopolymers) in terrigenous fossils and the chemical and structural changes that they undergo during fossilization are discussed over three critical stratigraphic levels or “time slices.” The youngest of these is the archeological record (e.g., <10 k.y. B.P.), when organic matter from living organisms undergoes the preliminary stages of fossilization (certain classes of biomolecule are selectively preserved while others undergo rapid degradation). The second time slice is the Tertiary. Well-preserved fossils of this age retain diagenetically modified biomarkers and biopolymers for which a product-precursor relationship with the original biological materials can still be identified. The final time slice is the Carboniferous. Organic material of this age has generally undergone such extensive diagenetic degradation that only the most resistant biopolymers remain and these have undergone substantial modification. Trends through time in the taphonomy and utility of ancient biomolecules in terrigenous fossils affect their potential for studies that involve chemosystematic and environmental data.