The extent to which perceived patterns of evolution are affected by the use of single characters versus overall morphology or measured versus counted and coded characters is explored empirically, employing multiple-character data from closely spaced sequential populations of the Neogene bryozoan Metrarabdotos. In all nine species examined, the pattern of evolution in overall morphology revealed by discriminant analysis is so highly punctuated that changes within species do not vary significantly from zero. Among the 46 single characters in the nine species, a few statistically significant temporal trends do occur, as apparent departures from the static pattern in overall morphology. However, these exceptions are so poorly related to the morphology that distinguishes inferred ancestor and descendant species from each other, and are so few in number, that they can be interpreted as chance variation from a general condition of no change within species. There is no difference in behavior between measured characters and counted or coded ones, in part because the distinction between the two types of characters can be artificial in bryozoans and other modular organisms. The results suggest that interpretation of single-character changes, in isolation rather than as statistical samples of the change in overall morphology, should be made with caution.

Arm autotomy was induced in a living specimen of Metacrinus rotundus (Echinodermata: Crinoidea). An arm was autotomized at a ligamentary articulation known as a cryptosyzygy, following incision by scissors distal to the break point. Although sessile stalked crinoids cannot entirely escape from a predatory attack by arm autotomy and they do not have an active defense, arm autotomy at cryptosyzygies reduces damage and arm loss by effective distribution, and by minimizing trauma and facilitating subsequent regeneration.

The paradigmatic distribution of cryptosyzygies in which arm loss is set at a minimum, compared with the actual distribution, shows that these two patterns are similar and that actual specimens successfully reduce arm loss by the effective distribution of cryptosyzygies. The crinoid branching pattern also affects arm loss, and two different paradigms are discussed: anti-predatory and harvesting. Arm branching patterns of various isocrinids have tended toward the anti-predatory configuration from the Jurassic to the Recent, suggesting that the isocrinids have coped with increased predation. Shallow-water comatulids generally adopt the anti-predatory paradigm in their branching pattern, whereas many deep-water, stalked crinoids adopt a harvesting paradigm, reflecting that shallow-water comatulids receive more predatory attacks than do deep-water crinoids.

Digitate shell morphologies have evolved repeatedly in planktonic foraminifera throughout the Cretaceous and Cenozoic. Digitate species are usually rare in fossil and modern assemblages but show increased abundance and diversity at times during the Cretaceous and middle Eocene. In this paper we discuss the morphology and stratigraphic distribution of digitate planktonic foraminifera and establish the isotopic depth ecology of fossil ones to draw parallels with modern counterparts. δ18O and δ13C values of six extinct and two modern digitate species, from six time slices (Cenomanian, Turonian, Eocene, Miocene, Pleistocene and Holocene) have similar isotopic depth ecologies, consistently registering the most negative δ13C and usually the most positive δ18O compared to coexisting species. These results indicate a similar deep, subthermocline (é150 m) habitat, characterized by lower temperatures, reduced oxygen, and enrichment of dissolved inorganic carbon. This is consistent with water-column plankton studies that provide insight into the depth preferences of the three modern digitate species; in over 70% of observations digitates occurred in nets below 150 m, and down to 2000 m. The correlation between digitate species and subsurface habitats across multiple epochs suggests that elongated chambers were advantageous for survival in a deep mesopelagic habitat, where food is usually scarce. Increased abundance and diversity of digitates in association with some early and mid-Cretaceous oceanic anoxic events, in middle Eocene regions of coastal and equatorial upwelling, and occasionally in some modern upwelling regions, suggests an additional link with episodes of enhanced ocean productivity associated with expansion of the oxygen minimum zone (OMZ). We suggest that the primary function of digitate chambers was as a feeding specialization that increased effective shell size and food gathering efficiency, for survival in a usually food-poor environment, close to the OMZ. Episodes of increased digitate abundance and diversity indicate expansion of the deep-water ecologic opportunity under conditions that were unfavorable to other planktonic species. Our results provide evidence of iterative evolution reflecting common functional constraints on planktonic foraminifera shell morphology within similar subsurface habitats. They also highlight the potential of digitate species to act as indicators of deep watermasses, especially where there was expansion of the OMZ.

Six compilations of fossil tetrapod families, spanning 100 years, each contain a broadly similar diversity pattern since the Upper Devonian. Comparison of four recent data bases, one of which is derived from a strict cladistic treatment, reveals widespread taxonomic and stratigraphic inaccuracies in three earlier data bases. Improvement of our interpretation of the tetrapod fossil record will come through continued taxonomic and stratigraphic revision as well as discovery of new fossils.

Addition of the recently discovered fossil Nalacetus to a phylogenetic analysis of basicranial, cranial, dental, postcranial, and soft morphological characters reveals that it is the most basal cetacean, and that mesonychians form the monophyletic sister group to Cetacea. The molars of Nalacetus elucidate transformations in dental morphology that occurred early in the cetacean radiation and clarify certain derived differences in molar cusp position between cetaceans and the extinct clade, Mesonychia, hypothesized to be their sister taxon. Nalacetus and other archaic cetaceans share derived vertically elongate shearing facets on the lower molars. Applying the Extant Phylogenetic Bracket, we advance the hypothesis that these facets are an osteological correlate of aquatic predation. Our functional interpretation of this character and its distribution within Cetacea indicates that a behavioral change in tooth use characterized the origin of the clade. Comparison of the transformation of this dental character with that of the cetacean pelvis indicates that a change in tooth use (feeding behavior) occurred before loss of the ability to engage in terrestrial locomotion.

The most parsimonious phylogenetic hypothesis presented here has a significant fit with the stratigraphic record as determined by the Manhattan Stratigraphic Measure, which is corroborated by retention indices of stratigraphic data. Ghost lineages necessitated by the phylogenetic hypothesis extend the stratigraphic range of Cetacea into the middle Paleocene (Torrejonian), ten million years earlier than the oldest cetacean fossil currently known. Primitive features of Nalacetus, the large number of synapomorphies diagnosing Cetacea, and the implied ghost lineage suggest that the early cetacean radiation was much more extensive than has been previously recognized.

Recent research has corroborated the long-held view that the diversity of genera within benthic marine communities has increased from the Paleozoic to the Cenozoic as much as three- to fourfold, after mitigating for such biasing influences as secular variation in time-averaging and environmental coverage. However, these efforts have not accounted for the considerable increase in the availability of unlithified fossiliferous sediments in strata of late Mesozoic and Cenozoic age. Analyses presented here on the Cenozoic fossil record of New Zealand demonstrate that unlithified sediments not only increase the amount of fossil material and hence the observed diversity therein, but they also preserve a pool of taxa that is compositionally distinct from lithified sediments. The implication is that a large component of the difference in estimates of within-community diversity between Paleozoic and Cenozoic assemblages may relate to the increased availability of unlithified sediments in the Cenozoic.

The macroevolutionary processes underlying large-scale biotic replacements are still poorly understood. Opinion remains divided regarding the roles of mass extinction, biotic interaction, and environmental perturbations in these replacement events. Previous attempts to test replacement hypotheses have largely focused on taxonomic diversity patterns. Taxonomic data alone, however, provide little insight about ecological interactions and hence other approaches are needed to understand mechanics of biotic replacements. Here I propose a conceptual model of replacement based on predation-mediated biotic interactions, and attempt a test using analysis of the Cenozoic replacement of the gastropod family Aporrhaidae by a closely related group, the Strombidae.

Taxonomic, morphologic, and geographic data analyzed in this study all suggest a replacement of aporrhaids by strombids following the end-Cretaceous mass extinction. While most of the taxonomic replacement was associated with a mass extinction, some replacement also occurred during background times and was mediated by higher origination rates in strombids rather than by higher extinction rates in aporrhaids. Morphologically, the replacement was largely confined to the portion of the morphospace unaffected by the end-Cretaceous extinction. At a global scale, the geographic overlap between the two groups declined through the Cenozoic, reflecting increasing restriction of aporrhaids to colder, temperate waters while strombids flourished in the tropics. However, at a finer geographic scale a more mosaic pattern of replacement is evident and coincides with Eocene and Oligocene climatic fluctuations.

The results of this study suggest that mass extinction, long-term biotic interaction, and environmental change can all play significant roles in biotic replacements. Since the relative importance of each factor would vary from one event to another, an understanding of the general nature of large-scale biotic replacements requires a knowledge of the relative intensities of each of these processes.

This study examines the morphology and function of hindlimb plumage in Archaeopteryx lithographica. Feathers cover the legs of the Berlin specimen, extending from the cranial surface of the tibia and the caudal margins of both tibia and femur. These feathers exhibit features of flight feathers rather than contour feathers, including vane asymmetry, curved shafts, and a self-stabilizing overlap pattern. Many of these features facilitate lift generation in the wings and tail of birds, suggesting that the hindlimbs acted as airfoils. A new reconstruction of Archaeopteryx is presented, in which the hindlimbs form approximately 12% of total airfoil area. Depending upon their orientation, the hindlimbs could have reduced stall speed by up to 6% and turning radius by up to 12%. Presence of the “four-winged” planform in both Archaeopteryx and basal Dromaeosauridae indicates that their common ancestor used fore- and hindlimbs to generate lift. This finding suggests that arboreal parachuting and gliding preceded the evolution of avian flight.

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

The shells of gastropods and cephalopods grow by production of new material, by the mantle, at the lip of the shell. I derive a model of shell form that allows us to describe the morphology of a shell in terms of independently definable biological parameters. These are (1) the relative rates of shell production at different points around the aperture, (2) the total amount of shell produced per time interval, (3) the growth rate of the aperture, (4) aperture shape, and (5) the orientation of the animal within the shell. Describing shell form in these terms allows us to see what biological changes must occur in development in order to change one shell morphology into another and what constraints are associated with particular morphological transformations. The model shows that it is developmentally easy to derive a slightly coiled limpet shell from that of a high-spired ancestor, but difficult to take the next step to a fully conical limpet. Many, if not most, real gastropod shells are not conical but rather have a convex or concave profile. I show that these forms result from a decoupling of shell production rates from the growth rate of the animal within the shell. The model also shows how truly different forms, such as vermetid snails and heteromorph ammonites, escaped the confines of spiral growth–sometimes by rotating the body within the shell, and sometimes by taking up a growth strategy that does not constrain them to coil. This model is compatible with shell morphometric models that have been widely discussed in the literature but strives toward a different goal: understanding the relationships between the various biological processes involved in shell development.

Living echinoids provide a structural analogue to the rigid thecae of many extinct echinoderms and are here used to examine some consequences of the increased rigidity and decreased number of plates which occurred in several lineages of echinoderms. A special feature of the echinoids is the large spines which protect the echinoid tests from failure under impact. The spines absorb energy or spread loads because the spines break or the tissue connecting the spine to the test is stretched or torn. Spines that absorb impacts might be superfluous in echinoderms encased in a rigid test or theca if they have flexible stems or are in low energy habitats, but spines should be useful to stemless thecate echinoderms on hard substrata. In the absence of spines, a flexible theca of overlapping plates might be superior to a rigid theca. Greater rigidity does not necessarily provide greater protection.

Tracts of podial pores are sites of weakness in all tests examined [Eucidaris tribuloides (Lamarck), Echinometra viridis Agassiz, Diadema antillarum Philippi, Lytechinus variegatus (Lamarck), Tripneustes ventricosus (Lamarck)]. Therefore arrangement of pores affects strength of rigid thecae. Sutures are stronger than plates in most species examined, but cracks cross sutures, and tests of E. tribuloides frequently break along sutures. It is not yet proven that sutures increase toughness of rigid tests by relieving stress on plates or stopping cracks. If resisting failure under impact is important, then preconditions for acquiring rigidity are life in a low energy habitat or accessory structures which absorb or spread impacts.

The pattern of differential species longevities among five Osagean–Meramecian crinoid clades is analyzed for its evolutionary significance. Differences in mean species longevity between clades may have resulted from species sorting based on eurytopy (niche breadth). In order to test the relationship between longevity and eurytopy it was first necessary to recognize generalists (eurytopes) vs. specialists (stenotopes) objectively. Three different approaches were used: (1) the “Eurytopy Index” (EI), which is a measure of mean number of facies per species; (2) analysis of crinoid functional morphology; and (3) use of canonical discriminant analysis to analyze species distributions between facies in order to separate generalists from specialists. Mean species longevity for each clade was evaluated by four different approaches: (1) rarefaction was used to control for differences in sample size, including both species richness and number of occurrences, between clades; (2) potential facies control of species longevity was evaluated by a bootstrap that compared the observed data to a null model where species longevity was limited only by the actual occurrences of each species known facies through time; (3) uniformity of clade species richness through time was evaluated by the “Index of Uniformity for Species Richness” based on the standard deviation of clade species richness across the time intervals; and (4) potential species range truncations were evaluated by a biostratigraphic gap analysis based on the binomial distribution.

The general order of increasing longevity and eurytopy is (from least to most): flexibles, advanced cladids, camerates, disparids, and primitive cladids. In general the pinnulate crinoids (advanced cladids and camerates) were specialists with lower mean species longevity, and the non-pinnulate crinoids (disparids and primitive cladids) were generalists with higher mean species longevity. Pinnulate crinoids were specialized for feeding in high-energy currents and, thus, were limited in their facies distribution and presumably more extinction-prone. The non-pinnulates could feed in both low- and high-energy currents and, thus, were less limited in their facies distribution and presumably less extinction-prone. The flexibles were the exception in that they were non-pinnulate but had the lowest mean species longevity, apparently because they were specialized for deeper-water clastic environments.

On average, generalist clades have mean species longevities that at a minimum are up to 45% (≈1.0 ± 0.7 m.y.) longer than specialist clades. However, greater mean species longevity did not necessarily confer long-term advantages to a clade. The specialist advanced cladids became the dominant crinoid clade of the late Paleozoic and gave rise to the articulate crinoids of the post-Paleozoic. This may have resulted from the more rapid species turnover of stenotopes creating adaptive evolutionary novelties for their clade. Alternatively, it could simply be the result of stochastic processes.

The finer subdivision of niche space by specialists has led previous workers to predict that specialist clades should have higher species richness than generalist clades. The present study supports this prediction.

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

A model for the paleobiogeographic history of South American cricetine rodents is proposed based on new and/or recently published fossil, geological, paleobotanical and radioisotope data. Cricetine rodents of the tribe Sigmodontini evolved in North America before 7.0 Myr BP. They got to South America by waif dispersal across the Bolivar Trough marine barrier from Central America during a world wide drop in sea level (the “Messinian Low”) between 7.0 and 5.0 Myr BP. The basal stock was probably a sylvan (forest) form, from which evolved pastoral (grazing) forms in the savanna-grassland area of Venezuela, Colombia and the Guianas. The pastoral forms in the northern savanna-grassland area were restricted there until about 3.5 Myr BP. At that time there occurred the first glaciation in South America and consonant with glacial advance was a retraction of forest habitats and an expansion of savanna-grassland habitats. At that time the pastoral forms were able to disperse southward through a savanna-grassland corridor along the eastern foothills of the Andes and spread throughout the previously disjunct savanna-grasslands of Bolivia and Argentina. Cricetines are first recorded as fossil in the Monte Hermoso Fm. of Argentina which is about 3.5 Myr BP in age. The Panamanian land bridge came into existence about 3.0 Myr BP as indicated by the beginning of a major interchange of terrestrial faunas between the Americas, which was well underway by 2.7 Myr BP.

Unfossiliferous marine sedimentary rocks of Phanerozoic age are known to all field-oriented paleontologists. These troublesome units are often encountered in the field, perhaps cursed roundly for a moment or two, and usually shrugged off in pursuit of the next fossiliferous interval. Paleontologists tend not to discuss barren units, and they rarely publish on the absence of a fauna from what appears to be unaltered marine rock. But aren't barren marine sediments revealing something important about their paleoenvironment and possibly about the paleoenvironments of conformably adjacent fossil-bearing units? Shouldn't paleontologists be just as interested in knowing the locations and ages of unfossiliferous sediments as they are fossiliferous strata?

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

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

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

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

The carbon stable isotope (δ13C) composition of the calcitic tests of planktonic foraminifera has an important role as a geochemical tracer of ocean carbon system changes associated with the Cretaceous/Paleogene (K/Pg) mass extinction event and its aftermath. Questions remain, however, about the extent of δ13C isotopic disequilibrium effects and the impact of depth habitat evolution on test calcite δ13C among rapidly evolving Paleocene species, and the influence this has on reconstructed surface-to-deep ocean dissolved inorganic carbon (DIC) gradients. A synthesis of new and existing multispecies data, on the relationship between δ13C and δ18O and test size, sheds light on these issues. Results suggest that early Paleocene species quickly radiated into a range of depths habitats in a thermally stratified water column. Negative δ18O gradients with increasing test size in some species of Praemurica suggest either ontogenetic or ecotypic dependence on calcification temperature that may reflect depth/light controlled variability in symbiont photosynthetic activity. The pattern of positive δ13C test-size correlations allows us to (1) identify metabolic disequilibrium δ13C effects in small foraminifera tests, as occur in the immediate aftermath of the K/Pg event, (2) constrain the timing of evolution of foraminiferal photosymbiosis to 63.5 Ma, ∼0.9 Myr earlier than previously suggested, and (3) identify the apparent loss of symbiosis in a late-ranging morphotype of Praemurica. These findings have implications for interpreting δ13C DIC gradients at a resolution appropriate for incoming highly resolved K/Pg core records.

The Caune de l'Arago Cave (southern France) has yielded one of the best preserved and best documented sedimentary successions of the European Middle Pleistocene (Oxygen Isotopic Stages 14 to 12). Herbivorous ungulates (horse, reindeer, red deer, fallow deer, bison, musk ox, argali, and tahr) are well represented in the three major stratigraphic units CM1, CM2, and CM3. CM1 and CM3 correspond to cold and dry climate and CM2 represents temperate and humid environmental conditions. Dental microwear and mesowear analyses were performed for the ungulates from CM1–3 to test whether these methods of dental wear evaluation were suitable for detecting climate-driven changes in the dietary resources of the Arago ungulate community. We found that both dental mesowear and microwear indicate dietary traits and their relationship to climatic conditions as reflected by vegetation cover and community structure. In all units, even if some species seem to share habitats or resources, it appears that the overlap in their feeding ecology is very low. The CM1 and CM3 units, where pollen analysis indicates that the climate was cold and dry, show the lowest diversity in dietary traits. The CM2, where climate is known to be more temperate and humid, the spectrum of dietary traits is large—grazers, browsers, and mixed feeders are present.

We use two approaches to test hypotheses regarding function in a group of extinct mammals (Family Mesotheriidae, Order Notoungulata) that lack any close extant relatives: a principle-derived paradigm method and empirically derived analog method. Metric and discrete morphological traits of mesotheriid postcranial elements are found to be consistent with the morphology predicted by a modified version of Hildebrand's paradigm for scratch diggers. Ratios of in-force to out-force lever arms based on skeletal elements indicate that the mesotheriids examined had limbs modified for high out-forces (i.e., they were “low geared”), consistent with the digging hypothesis. Other mesotheriid characters, such as cleft ungual phalanges, a curved olecranon, and a highly modified pelvis (with extra vertebrae incorporated into the sacrum and fusion between the ischium and the axial skeleton) are regarded as being functionally significant for digging and also occur in a variety of extant diggers. Analog methods indicate that mesotheriids share numerous traits common to a variety of extant diggers. Principal component analyses of postcranial elements indicate that mesotheriids consistently share morphometric space with larger extant fossorial mammals: aardvark, anteaters, wombats, and badger. Likewise, discriminant function analyses categorized mesotheriids as fossorial, though imperfectly analogous to the extant diggers analyzed. Thus, both theory-driven and empirically derived methods of estimating function in these extinct taxa support a digging hypothesis for the mesotheriids examined. Adaptations for digging in both the forelimb and sacropelvic functional complexes of mesotheriids provide independent support for the fossorial hypothesis.