Coiled cephalopods constitute a major part of the Paleozoic nekton. They emerged in the Early Ordovician but nearly vanished in the Silurian. The Emsian appearance of ammonoids started a story of evolutionary success of coiled cephalopods, which lasted until the end-Permian extinction event. This story is investigated by using a taxonomic database of 1346 species of 253 genera of coiled nautiloids and 1114 genera of ammonoids. The per capita sampling diversities, the Van Valen metrics of origination and extinction, and the probabilities of origination and extinction were calculated at stage intervals. The outcome of these estimations largely reflects the known biotic events of the Paleozoic. The polyphyletic, iterative appearance of coiled cephalopods within this time frame is interpreted to be a process of adaptation to shell-crushing predatory pressure. The evolution of the diversity of coiled nautiloids and ammonoids is strongly correlated within the time intervals. Once established, assemblages of coiled cephalopods are related to changes in sea level. The general trends of decreasing mean (or background) origination and extinction rates during the Paleozoic are interpreted to reflect a successive stabilization of the coiled cephalopod assemblages. Different reproduction strategies in ammonoids and nautiloids apparently resulted in different modes of competition and morphological trends. Significant morphological trends toward a stronger ornamentation and a centrally positioned siphuncle characterize the evolution of Paleozoic nautiloids.

More than 300 coots (Fulica americana) became frozen in Spring Lake, Tazewell County, Illinois, on December 1, 1985. This catastrophic event permitted 8 weeks of taphonomic observations, which showed that ice forms a stable substrate which permits terrestrial taphonomic processes to be imprinted on lacustrine deposits. Bird and mammal scavengers attacked coot carcasses in different manners, resulting in distinct disarticulation sequences. Bird scavengers preferentially fed on the head, neck, and breast-wing complex, causing early disarticulation of bones in these areas, late loss of hindlimb joints, and minimal bone damage. In contrast, mammal scavengers concentrated their attention on the hindlimb and tail region, resulting in bone breakage and early disarticulation of these body parts, but late disarticulation of the breast-wing complex. These data demonstrate that scavenger-specific feeding behaviors significantly influence disarticulation patterns early in assemblage formation, while anatomy may exert increasingly greater influence on disarticulation patterns as carasses become less attractive to scavengers. Finally, because taphonomic processes change in intensity and type through time, bone frequency and modification patterns will vary according to the time at which the patterns are arrested by burial. Thus, bone frequency and modification patterns should provide an index to the relative importance of specific biotic agents and of anatomy in fossil disarticulation patterns as well as an estimate of the time between death and burial.

Shells of 375 sexed specimens of Nautilus cf. N. pompilius trapped at 300–700 ft (90–215 m) depth off Mutremdiu Point and Ngemelis Island, Palau, exhibit distinctive dimorphism in shell proportion and overall size. Of 221 mature animals, males have larger shells (mean diameter = 209 mm), a broader aperture (mean maximum width = 99 mm; mean width at apertural salient = 92 mm) and greater weight (mean shell + body weight = 1426 gm). Female shells are smaller (mean diameter = 198 mm), narrower (mean maximum width = 91 mm; mean apertural salient width = 81 mm) and the overall weight is less (mean = 1157 gm). Dimorphic differences are not apparent in young shells (less than 180 mm diameter), but develop during the final ½ to ¼ whorl. This is accompanied by development of the spadix in males, indicating that shell dimorphism reflects sexual maturity. Of 375 animals trapped, 28% were female; no depth segregation was apparent. Traditional identification of larger fossil dimorphic cephalopods as females and smaller forms as males is the reverse of observed dimorphism in Nautilus and should be discontinued in favor of macroconch and microconch designations.

Sabertooth members of the Felidae, Nimravidae, and Barbourofelidae are well-known for their elongated saber-shaped canines. However, within these groups, there is a wide range of independently derived tooth shapes and lengths, including dirk-tooth and scimitar-tooth morphs. In conjunction with the saberteeth, forelimbs were also used to subdue prey. Thus, there may be a functional link between canine shape and forelimb morphology. Because there are no living sabertooth forms for comparison, extant felids make a good proxy for examining the morphology of these extinct organisms. Here, I examine the forelimb morphology of different sabertooth groups from across North America; I address whether forelimb morphologies are associated with tooth morphologies, and whether these associated tooth and forelimb morphologies are convergent among different families. To answer these questions, I analyzed six functional indices of the forelimbs and two canine characters for 13 species of sabertooth predators and 15 extant felid species. Results indicate that sabertooth morphs with longer, thinner canines show more robust limb proportions. These patterns were convergent among sabertooth felids, nimravids, and barbourofelids, and indicate a positive functional relationship between saber elongation and increased forelimb robustness. This suggests that sabertooth carnivorans demonstrated niche partitioning of predation strategies according to canine shape and corresponding forelimb morphology.

Relative abundance data are of primary importance in paleoecology, but it is not always obvious how they should be interpreted. Because relative abundance is expressed as a proportion of the total sample, change in the abundance of one group necessarily changes the relative abundance of all groups in the sample. There are two possible interpretations for a trend in the relative abundance of a taxon: an “active” scenario in which the trend reflects change in the population density of the group itself, or a “passive” scenario in which the change is driven by population changes in other taxa. To discriminate between these scenarios it is necessary to collect absolute abundance data (abundance expressed as a function of sample area or volume).

We examine both absolute and relative abundance trends through a major paleoecological transition: the shift from trilobite-dominated to brachiopod-dominated paleocommunities in shallow marine carbonates spanning the Lower/Middle Ordovician boundary in western Utah and eastern Nevada. We sampled 61 carbonate mudstone and wackestone beds from the upper Ibex Series (Lower Ordovician) and lower Whiterock Series (Middle Ordovician) at three sections that span the boundary. All samples come from the shallow subtidal Bathyurid trilobite biofacies. Samples were broken into small pieces, and all skeletal fragments >2 mm were identified to the finest possible taxonomic level. Consistent with previous work on this interval, the relative abundance of trilobites declines sharply across the boundary, while the relative abundance of brachiopods increases. Absolute abundance data indicate that the decline in trilobite abundance is genuine and not an artifact of normalization. The trend is not easily explained by sampling bias, facies distribution, taphonomic regime, or sedimentation style.

The dramatic shift in abundance contrasts with relatively minor changes in relative genus richness across the boundary. This is partly ascribable to differences in the relative abundance structure of trilobite faunas. Though comparable numbers of trilobite and brachiopod genera occur above and below the boundary, the trilobite fauna from the upper Ibex Series has lower evenness then the lower Whiterock Series fauna. Hence sampled trilobite richness is high in the lower Whiterock despite the small number of specimens. This highlights the importance of collecting abundance data. Although these data suggest that in at least some cases richness and abundance patterns are not closely coupled, more robust richness data are necessary to confirm this conclusion.

Geographic range appears to be an important aspect of the biology of species, but ranges cannot be unambiguously determined from the fossil record: ancient species can rarely be traced to the full extent of their original geographic ranges, and some ancient provinces are far better characterized than others. Here we test the degree to which paleogeographic range data for species within ancient provinces can be used to estimate the relative magnitudes of total geographic ranges, focusing on the 212 living bivalve species of the Oregonian Province, a north-south (N-S) province of the northeastern Pacific shelf.

We break the total range of each species into its within-province and extraprovincial components. Nonparametric rank correlation tests and simple linear regressions yield highly significant correlations, both between within-province and total ranges (which are not independent variables) and between within-province and extraprovincial ranges. The strongest correlations are obtained when north-ranging species are excluded. These north-ranging species are thermally tolerant and have access to broad east-west (E-W) provinces, thus weakening the relation between their within-province and total ranges. On a coarser scale, we also found that the extraprovincial ranges of species with small within-province ranges (<650 km, ca. 25% of focal province) are significantly less than those for species with large within-province ranges (> 1950 km, ca. 75% of focal province). The significant correlations of within-province range with both total and extraprovincial ranges hold for species with within-province ranges >650 km, so caution is needed regarding species that extend into a focal province only a short distance from its edges. We conclude that within-province geographic range can generally be used to rank species by total geographic range and thus is an adequate proxy in most comparative studies of the paleobiologic consequences of range magnitudes.

Mammalian species often exhibit clinal geographic variation in body size: individuals tend to be larger in areas with lower mean annual temperature. Climatic change involving increasing or decreasing mean annual temperature may cause clines to shift geographically, resulting in a phenotypic shift at all affected locales within a species' range. I assess the potential of shifting geographic clines to produce morphological trends in the fossil record. Five extant North American mammalian species (Didelphis virginiana, Mephitis mephitis, Odocoileus virginianus, Scalopus aquaticus, and Sciurus carolinensis) are examined to quantify size change along latitudinal clines and to estimate the geographic range and temperature difference commonly associated with a given difference in body size. Relative to body size, the observed size range of skeletal characters within each of these five species is comparable to that seen in a much larger sample of North American mammals. Thus patterns of variation documented for the five species may be used to assess the likelihood of dine translocation as an explanation of size change in the mammalian fossil record. As a case study, I examine three lineages from the Early Eocene of the Bighorn Basin, Wyoming. I determine that size change in these chronoclines represents evolutionary change and is not merely the result of shifting geographic clines.

Open-ended tubes were found extending inward from the inner surfaces of brachial valves of three different species of Emsian spiriferid brachiopods. It is believed that these tubes were formed by the outer epithelium of the brachiopods, precipitating shell material around a small commensal filter feeding organism living in the outer mantle cavity. The position of the open ends of these tubes, the business end of the filter feeding organism, in every case discovered is between the bases of the two spiralia. This lends support to the suggestion of Ager (1961) and Wallace and Ager (1966) that spiriferid brachiopods had median inhalant and lateral exhalant feeding currents. These structures are named Burrinjuckia spiriferidophilia n.gen. n.sp. They are trace fossils of indeterminate taxonomic position.

Did organisms diversify in different ways on land and in the marine realm over the Phanerozoic, or do the different diversification curves of continental and marine organisms reflect primarily methodological artifacts? To answer this question, a quantitative assessment of the completeness of the global continental fossil record is indispensable. We used comparisons between continental and marine fossil diversity and between past and present-day patterns of continental diversity to assess the absolute and relative completeness of the continental fossil record. Collector's curves of the number of described families over the past 200 years suggest that the global continental fossil record, and even that of European and North American tetrapods, is still highly incomplete. Nevertheless, relative proportions of major continental and marine taxa, patterns of tetrapod endemism, and familial durations suggest that the family-level continental fossil record is reasonably representative. We found that, although continental fossil richness is correlated with the amount of terrestrial clastic sediment available for sampling, the exponential diversification curve of continental metazoans is unlikely to be an artifact of this rock bias. Diversification of the continental fauna appears to have been essentially exponential since the Devonian, with little evidence of major extinction events.

In a hyperbaric chamber, a living mature specimen of Nautilus pompilius withstood a hydrostatic pressure of 8.05 MPa (80.5 kg/cm2) equivalent to 785 m deep in the sea. Thereafter it was killed instantly by implosion of the shell. Before implosion, the animal reacted physiologically to increasing pressure. Therefore, the depth of 785 m can be assigned the depth limit of N. pompilius. The result bears on critical interpretations on the paleoecology and paleobiology of extinct nautiloids and ammonoids with similar shells.

Cretaceous floras in Alaska, when compared to those at mid-latitudes, generally indicate later appearances in Alaska of major clades and major leaf morphologies. Compared to mid-latitude floras, Alaskan Late Cretaceous floras contain few major clades. The Alaskan clades diversified but at a low taxonomic level. Migrational pathways into high latitudes were probably along streams. Similar patterns characterized the Alaskan Tertiary, although some southward migrations of lineages occurred during the Neogene.

Review of other Arctic paleontological data from Ellesmere Island, previously used to suggest that the Arctic was a major center of origin during the Late Cretaceous, indicates that the ages of supposedly substantiating dinoflagellate floras were misinterpreted. When the dinoflagellate data are interpreted according to standard methodology, first occurrences of genera and species groups on Ellesmere are, like the Alaskan occurrences, later than first occurrences at middle latitudes.

Incremental stages of major evolutionary transitions within a single animal lineage are rarely observed in the fossil record. However, the extraordinarily complete sequence of well preserved material spanning the 27-Myr existence of the marine squamate subfamily Mosasaurinae provides a unique exception. By comparison with extant and extinct analogs, the tail morphology of four mosasaurine genera is examined, revealing a pattern of evolution that begins with the generalized varanoid anatomy and culminates in a high-aspect-ratio fin, similar to that of sharks. However, unlike the epicercal caudal fluke of selachians in which the tail bends dorsocaudally, derived mosasaurs develop a hypocercal tail with a ventrocaudal bend. Progressive caudal regionalization, reduced intervertebral mobility, increased tail depth due to a marked downturn of the posterior caudal segment, and the development of finlike paired appendages reveal a pattern of adaptation toward an optimized marine existence. This change in morphology reflects a transition from anguilliform or sub-carangiform locomotion to carangiform locomotion, and indicates a progressive shift from nearshore dwellers to pelagic cruisers—a change in foraging habitat independently corroborated by paleobiogeographic, stable isotope, osteohistological, and paleopathological data. Evolutionary patterns similar to those observed in mosasaurine mosasaurs are seen in other secondarily aquatically adapted amniotes, notably metriorhynchid crocodyliforms, cetaceans, and ichthyosaurs, and may be explained by developmental modularity governing the observed phenotypic expression.

Spectral algorithms have been shown to work well in a wide range of situations that involve the task of ordering. When applied to the localities of a set of European Neogene land mammal taxa, spectral ordering relies almost entirely on the most common genera, depends on connectivity more than on length of taxon lists, and is robust to noise from rarer and less connected taxa. The spectral coefficients for localities are highly correlated with known geochronological ages. Although elementary compared with more sophisticated biochronological tools, spectral ordering allows a fast and standardized way to generate biochronological ordering of localities when other information than faunal lists is lacking. Compared with the conventional mammal Neogene (MN) units, spectral ordering of localities appears to lack distinct temporal boundaries in taxon content and render a much lower count of Lazarus events. If, as seems to be the case, biochronology depends mainly on the most common taxa and if evolutionary change is also most clearly reflected in them, then the main evolutionary patterns should be detectable at a modest level of sampling.

If last appearances of marine animal genera are taken as reasonable proxies for true extinctions, then there is appreciable global extinction in every stage of the Phanerozoic. If, instead, backsmearing of extinctions by incomplete sampling is explicitly taken into consideration, a different view of extinction emerges, in which the pattern of extinction is much more volatile and in which quiescent time spans—with little or no global extinction for several million years—are punctuated by major extinction events that are even more extreme than is generally thought. Independent support for this alternative view comes from analysis of genus occurrence data in the Paleobiology Database, which agrees with previous estimates of sampling probability and implies that offsets between extinction and last appearance of one or more stages are quite probable.

The ammonoid order Prolecanitida constitutes a relatively small (43 genera, ~250 species) but long-ranging lineage (Lower Carboniferous—Triassic, ~108 m.y.), which narrowly survived the P/Tr extinctions and provided the stock from which were derived all later Mesozoic ammonoids. Prolecanitids were a minority among Late Paleozoic ammonoids, which were dominated by the Goniatitida, and showed many features that set them far apart from their contemporaries, including (1) long-term, gradual changes in shell geometry (W-D-S); (2) the most strongly constrained morphospace of any Paleozoic ammonids examined to date; (3) an eight-fold increase in mean suture complexity (three times that of Pennsylvanian goniatitids); (4) high correlations between shell geometry, shell and septal thickness, and suture complexity; (5) short body chambers and, as a consequence, high aperture orientations; (6) indications that cameral liquid may have been used for buoyancy control; and (7) a genus longevity that averaged 14.7 m.y. compared with 5.7 m.y. in Upper Carboniferous goniatitids, and that appears to have been unrelated to suture complexity. Prolecanitids showed a pervasive tendency to increase suture complexity (in the clade as a whole as well as within subclades and in more than 90 percent of ancestor-descendant genera), thus arguing a case for a driven complexity trend. The uniqueness of the prolecanitids calls into question whether they and their Mesozoic descendants, ceratites and ammonites, were strictly analogous to Paleozoic goniatites.

The chambered shell of modern cephalopods functions as a buoyancy apparatus, allowing the animal to enter the water column without expending a large amount of energy to overcome its own weight. Indeed, the chambered shell is largely considered a key adaptation that allowed the earliest cephalopods to leave the ocean floor and enter the water column. It has been argued by some, however, that the iconic chambered shell of Paleozoic and Mesozoic ammonoids did not provide a sufficiently buoyant force to compensate for the weight of the entire animal, thus restricting ammonoids to a largely benthic lifestyle reminiscent of some octopods. Here we develop a technique using high-resolution computed tomography to quantify the buoyant properties of chambered shells without reducing the shell to ideal spirals or eliminating inherent biological variability by using mathematical models that characterize past work in this area. This technique has been tested on Nautilus pompilius and is now extended to the extant deep-sea squid Spirula spirula and the Jurassic ammonite Cadoceras sp. hatchling. Cadoceras is found to have possessed near-neutral to positive buoyancy if hatched when the shell possessed between three and five chambers. However, we show that the animal could also overcome degrees of negative buoyancy through swimming, similar to the paralarvae of modern squids. These calculations challenge past inferences of benthic life habits based solely on calculations of negative buoyancy. The calculated buoyancy of Cadoceras supports the possibility of planktonic dispersal of ammonite hatchlings. This information is essential to understanding ammonoid ecology as well as biotic interactions and has implications for the interpretation of geochemical data gained from the isotopic analysis of the shell.

Throughout the late Quaternary, the Sahul (Pleistocene Australia–New Guinea) vertebrate fauna was dominated by a diversity of large mammals, birds, and reptiles, commonly referred to as megafauna. Since ca. 450–400Ka, approximately 88 species disappeared in Sahul, including kangaroos exceeding 200kg in size, wombat-like animals the size of hippopotamuses, flightless birds, and giant monitor lizards that were likely venomous. Ongoing debates over the primary cause of these extinctions have typically favored climate change or human activities. Improving our understanding of the population biology of extinct megafauna as more refined paleoenvironmental data sets become available will assist in identifying their potential vulnerabilities. Here, we apply a multiproxy approach to analyze fossil teeth from deposits dated to the middle and late Pleistocene at Cuddie Springs in southeastern Australia, assessing relative aridity via oxygen isotopes as well as vegetation and megafaunal diets using both carbon isotopes and dental microwear texture analyses. We report that the Cuddie Springs middle Pleistocene fauna was largely dominated by browsers, including consumers of C4 shrubs, but that by late Pleistocene times the C4 dietary component was markedly reduced. Our results suggest dietary restriction in more arid conditions. These dietary shifts are consistent with other independently derived isotopic data from eggshells and wombat teeth that also suggest a reduction in C4 vegetation after ~45 Ka in southeastern Australia, coincident with increasing aridification through the middle to late Pleistocene. Understanding the ecology of extinct species is important in clarifying the primary drivers of faunal extinction in Sahul. The results presented here highlight the potential impacts of aridification on marsupial megafauna. The trend to increasingly arid conditions through the middle to late Pleistocene (as identified in other paleoenvironmental records and now also observed, in part, in the Cuddie Springs sequence) may have stressed the most vulnerable animals, perhaps accelerating the decline of late Pleistocene megafauna in Australia.

Previous studies have shown that secular variation in the Mg/Ca ratio of seawater throughout the Phanerozoic would have subjected the aragonite-producing codiacean algae to at least three transitions between the low-Mg calcite (molar Mg/Ca <2) and aragonite + high-Mg calcite (molar Mg/Ca >2) nucleation fields in the oceans, since their origin in the Ordovician. These studies have asserted that major sediment production by codiacean algae in Recent tropical seas is permitted by the Mg/Ca ratio of modern seawater (molar Mg/Ca ∼5.2) remaining within the aragonitic/high-Mg calcite nucleation field (molar Mg/Ca >2). Here I present the results of experiments conducted to determine the effects of ambient Mg/Ca on the mineralogy, growth rate, primary productivity, calcification rate, and biomechanics of the codiacean alga Penicillus capitatus.

P. capitatus specimens were grown in three artificial seawater treatments that mimic ancient seawater of differing Mg/Ca ratios, corresponding to the low-Mg calcite nucleation field (molar Mg/Ca ∼1.0), a “boundary field” (molar Mg/Ca ∼2.5), and the aragonite + high-Mg calcite nucleation field (molar Mg/Ca ∼5.2). Significantly, P. capitatus specimens maintained a mostly aragonitic mineralogy in all three seawater treatments. However, linear growth rates, primary productivity, calcification, and thallus stiffness decreased with reductions in ambient Mg/Ca. That P. capitatus precipitates approximately three-quarters of its CaCO3 as aragonite in the seawater treatment that favors the inorganic precipitation of low-Mg calcite suggests that the alga dictates the precipitation of that polymorph, either by pumping cations to create an internal aragonite nucleation field (molar Mg/Ca >2) or by employing organic templates that specify the nucleation of the aragonite polymorph (Borowitzka 1984). However, the alga's precipitation of one-quarter of its CaCO3 as low-Mg calcite suggests that its mineralogical control is limited and can be partially overridden by the Mg/Ca ratio of ambient seawater. The reduced calcification of P. capitatus specimens grown in the low-Mg calcite and boundary nucleation fields is probably due to the inherent difficulty of precipitating aragonite from seawater which does not naturally support its nucleation. The decreased rates of linear growth and primary production are probably caused by reductions in CO2 available for photosynthesis due to the reduction in calcification (Borowitzka and Larkum 1977). The observed decrease in thallus stiffness is probably due to the reductions in calcification and primary productivity observed in P. capitatus specimens grown in the low-Mg calcite and boundary nucleation fields.

The present study suggests that aragonitic algae would have been handicapped in oceans characterized by Mg/Ca ratios that did not support their inherent mineralogy. Producing aragonite in seawater outside of the aragonite + high-Mg calcite nucleation field would probably have reduced the competitiveness of these algae, made them more susceptible to predation, and reduced their contribution to carbonate sedimentation. These findings support earlier assertions that the dominant ecological and sedimentological roles of codiacean algae in Recent tropical seas is permitted by a Mg/Ca ratio of seawater that supports the algae's aragonitic mineralogy during this time.

At present there are many animal phyla that contain only a few species. The existence of these small phyla can be used to test assumptions about speciation and extinction in multicellular animals.

We first model the number of species in a monophyletic clade with a birth and death process that assumes rates of speciation and extinction are constant. If no new phyla have evolved since the Cambrian and speciation and extinction rates for minor phyla are similar to or greater than those estimated from fossils, then our model shows that the probabilities of minor phyla surviving to the present are small. Random variation in extinction and speciation rates does not improve the chances for persistence. If speciation rates exceed extinction rates at the initial radiation of the clade, but before the clade becomes large, speciation rates come to equal extinction rates and both are low, persistence from before the Ordovician up to the present becomes likely. These models show that if speciation and extinction rates are independent of the number of species in a clade, then conditions before the Ordovician strongly influence today's distribution of species among taxa.

We also discuss a model in which speciation and extinction rates depend on the number of species in a clade. This alternative model can account for the persistence of phyla with few species to the present and predicts a short duration for phyla that did not exceed a threshold number of species.