Sexual dimorphism is documented in 35 articulated adult skeletons, 24 females, and 11 males, of the Miocene rhinoceros Teleoceras major from Ashfall Fossil Beds, Nebraska. Morphometric analysis of 51 cranial, mandibular, forelimb, and hindlimb characters reveals larger male mean values in 50 of the 51 measurements, of which 23 are significantly different (p ≤ 0.01). The most clearly dimorphic feature is the i2 diameter. The dimorphism evident in additional mandibular and cranial characters is conservative when compared with the dimorphism present in the fore- and hindlimbs. Non-overlapping male and female ranges are recorded for humerus length, radius length, radius proximal width, and femur length, with corresponding dimorphism ratios (DR = male ÷ female) of 1.11, 1.12, 1.11, and 1.10. Maximum male longbone lengths exceed minimum female lengths by an average of 24% (20–29%). Developmental maturity is apparently asynchronous in T. major, with fusion of longbone epiphyses delayed a minimum of two relative adult age classes in males. Significant sexual dimorphism is evident in the radius (DR = 1.34) and femur (DR = 1.19) cross-sectional areas. Estimates of body mass suggest a DR value between 1.13 and 1.23. The cranial, mandibular, and body-size dimorphism in T. major approaches that seen in the extant rhinoceroses Ceratotherium simum and Rhinoceros unicornis. However, the apparent herd structure and breeding-age sex ratio for the Ashfall herd suggests a behavioral ecology for T. major different from that of extant rhinoceroses. Teleoceras was likely a herding polygynous species ecologically more similar to extant Hippopotamus amphibius of Africa.

Atmospheric carbon dioxide is the raw material for the biosphere. Therefore, changes in the carbon isotopic composition of the atmosphere will influence the terrestrial δ13C signals we interpret. However, reconstructing the atmospheric δ13C value in the geologic past has proven challenging. Land plants sample the isotopic composition of CO2 during photosynthesis. We use a model of carbon isotopic fractionation during C3 photosynthesis, in combination with a meta–data set (519 measurements from 176 species), to show that the δ13C value of atmospheric CO2 can be reconstructed from the isotopic composition of plant tissue. Over a range of pCO2 (198–1300 ppmv), the δ13C value of plant tissue does not vary systematically with atmospheric carbon dioxide concentration. However, environmental factors, such as water stress, can influence the δ13C value of leaf tissue. These factors explained a relatively small portion of variation in the δ13C value of plant tissue in our data set and emerged strongly only when the carbon isotopic composition of the atmosphere was held constant. Members of the Poaceae differed in average δ13C value, but we observed no other differences correlated with plant life form (herbs, trees, shrubs). In contrast, over 90% of the variation the carbon isotopic composition of plant tissue was explained by variation in the δ13C value of the atmosphere under which it was fixed. We use a subset of our data spanning a geologically reasonable range of atmospheric δ13C values (−6.4‰ to −9.6‰) and excluding C3 Poaceae to develop an equation to reconstruct the δ13C value of atmospheric CO2 based on plant values. Reconstructing the δ13C value of atmospheric CO2 in geologic time will facilitate chemostratigraphic correlation in terrestrial sediments, calibrate pCO2 reconstructions based on soil carbonates offer a window into the physiology of ancient plants.

The environmental and biotic history of the late Quaternary represents a critical junction between ecology, global change studies, and pre-Quaternary paleobiology. Late Quaternary records indicate the modes and mechanisms of environmental variation and biotic responses at timescales of 101–104 years. Climatic changes of the late Quaternary have occurred continuously across a wide range of temporal scales, with the magnitude of change generally increasing with time span. Responses of terrestrial plant populations have ranged from tolerance in situ to moderate shifts in habitat to migration and/or extinction, depending on magnitudes and rates of environmental change. Species assemblages have been disaggregated and recombined, forming a changing array of vegetation patterns on the landscape. These patterns of change are characteristic of terrestrial plants and animals but may not be representative of all other life-forms or habitats. Complexity of response, particularly extent of species recombination, depends in part on the nature of the underlying environmental gradients and how they change through time. Environmental gradients in certain habitats may change in relatively simple fashion, allowing long-term persistence of species associations and spatial patterns. Consideration of late Quaternary climatic changes indicates that both the rate and magnitude of climatic changes anticipated for the coming century are unprecedented, presenting unique challenges to the biota of the planet.

Ceratopsid dinosaurs traditionally have been restored with sprawling forelimbs and were considered unable to run at high speeds. An alternative view restores the ceratopsids as rhinoceros-like with parasagittal forelimb kinematics and the ability to run faster than extant elephants. Several anatomical difficulties concerning the mounting of ceratopsid skeletons with nearly parasagittal forelimbs stem not from the forelimb itself, but from errors in rib and vertebral articulation. Matching a skeletal restoration to a probable ceratopsid trackway shows that the hands were placed directly beneath the glenoids, and that manual impressions were directed laterally, not medially as in sprawling reptiles. Pedal impressions in trackways are medial to the manual impressions, owing to the slightly averted elbow and to the asymmetrical distal femoral condyles, which directed the crus slightly medially. The limbs of ceratopsians of all sizes display substantial joint flexure, strongly indicating that the elephantine forelimb posture that has sometimes been suggested as the alternative to a sprawling posture is erroneous. The articular surfaces of uncrushed ceratopsian scapulocoracoids and forelimb joints confirm that the forelimb operated in a near-parasagittal plane with the elbows only slightly averted. The maximal running speed of even the largest ceratopsids is inferred to have significantly exceeded that of elephants and was probably broadly similar to that of rhinos.

A new sampling technique for fossil bone (coring with a 5/8″ bit) was used to sample longbones of all four sauropod genera from the Upper Jurassic Tendaguru beds of Tanzania for paleohistological study. Brachiosaurus and Barosaurus are represented by growth series of humeri and femora, while Dicraeosaurus could be sampled in fewer specimens and only one bone of Janenschia was available. Although all samples are dominated by fibrolamellar bone tissue, taxa can be distinguished by the degree and nature of bone remodeling and the presence and spacing of a peculiar kind of growth line (here termed “polish lines”). In addition, Barosaurus bone revealed two types of histology, tentatively interpreted as sexual morphs. The Tendaguru sauropods show a common growth pattern in which growth is determinate but sexual maturity is achieved well before maximum size is reached. For Brachiosaurus and Barosaurus, size at sexual maturity can be estimated and was reached at about 40% and 70% maximum size, respectively. Quantification of growth is possible in Janenschia using polish lines: the specimen studied reached sexual maturity at ≥11 years, attained maximum size at ≥26 years, and died at ≥38 years.

The excellent fossil record of the past few million years, combined with the overwhelming similarity of the biota to extant species, provides an outstanding opportunity for understanding paleoecological and macroevolutionary patterns and processes within a rigorous biological framework. Unfortunately, this potential has not been fully exploited because of lack of well-sampled time series and adequate statistical analysis. Nevertheless, four basic patterns appear to be of general significance. First, a major pulse of extinction occurred 1–2 m.y. ago in many ocean basins, more or less coincident with the intensification of glaciation in the Northern Hemisphere. Rates of origination also increased greatly but were more variable in magnitude and timing. The fine-scale correlation of these evolutionary events with changes in climate is poorly understood. Similar events probably occurred on land but have not been tested adequately. Second, rates of origination and extinction in the oceans waned after the pulse of extinction, especially during the past 1 m.y. Thus, most marine species originated long before the Pleistocene under very different environmental circumstances, suggesting that they are “exapted” rather than adapted to their present ecological circumstances. The same may be true for many terrestrial groups, but not for the mammals or fresh-water fishes that have continued to undergo speciation throughout the Pleistocene. Third, community membership of late Pleistocene coral reef communities was more stable than expected by chance. These are the only paleoecological data adequate to test hypotheses of community stability, so that we do not know whether community structure involving other taxa or environments typically reflects more than the collective behavior of individual species distributions. Regardless, the strong evidence for nearly universal exaptation of ecological characteristics argues strongly against ideas of coevolution of species in communities. Finally, ecological communities were profoundly altered by human activities long before modern ecological studies began. Holocene paleontological, archeological, and historical data constitute the only ecological baseline for “pristine” ecological communities before significant human disturbance. Holocene records should be much more extensively used as a baseline for Recent ecological studies and for conservation and management.

This paper documents a series of methodological innovations that are relevant to macroevolutionary studies. The new methods are applied to updated faunal and body mass data sets for North American fossil mammals, documenting several key trends across the late Cretaceous and Cenozoic. The methods are (1) A maximum likelihood formulation of appearance event ordination. The reformulated criterion involves generating a maximally likely hypothesized relative ordering of first and last appearances (i.e., an age range chart). The criterion takes faunal occurrences, stratigraphic relationships, and the sampling probability of individual genera and species into account. (2) A nonparametric temporal interpolation method called “shrink-wrapping” that makes it possible to employ the greatest possible number of tie points without violating monotonicity or allowing abrupt changes in slopes. The new calibration method is used in computing provisional definitions of boundaries among North American land mammal ages. (3) Additional methods for randomized subsampling of faunal lists, one weighting the number of lists that have been drawn by the sum of the square of the number of occurrences in each list, and one further modifying this approach to account for long-term changes in average local species richness. (4) Foote's new equations for instantaneous speciation and extinction rates. The equations are rederived and used to generate time series, confirm that logistic dynamics result from the diversity dependence of speciation but not extinction, and define the median duration of species (i.e., 2.6 m.y. for Eocene-Pleistocene mammals). (5) A method employing the G likelihood ratio statistic that is used to quantify the volatility of changes in the relative proportion of species falling in each of several major taxonomic groups. (6) Univariate measures of body mass distributions based on ordinary moment statistics (mean, standard deviation, skewness, kurtosis). These measures are favored over the method of cenogram analysis. Data are presented showing that even diverse individual fossil collections merely yield a noisy version of the same pattern seen in the overall continental data set. Peaks in speciation rates, extinction rates, proportional volatility, and shifts in body mass distributions occur at different times, suggesting that environmental perturbations do not have simple effects on the biota.

Paleobiology publishes original contributions dealing with any biological aspect of the history of life. Emphasis is placed on biological or paleobiological processes and patterns, including macroevolution, extinction, diversification, speciation, functional morphology, biogeography, phylogeny, paleoecology, molecular paleontology taphonomy, among others. Papers will typically interest readers from more than one specialty. Proposals for symposium volumes should be discussed in advance with the editors. Larger manuscripts on the same subjects as those normally covered in Paleobiology articles should be submitted to the Paleobiology Memoirs series.

Pelagic (open-ocean) species have enormous population sizes and broad, even global, distributions. These characteristics should damp rates of speciation in allopatric and vicariant evolutionary models since dispersal should swamp diverging populations and prevent divergence. Yet the fossil record suggests that rates of evolutionary turnover in pelagic organisms are often quite rapid, comparable to rates observed in much more highly fragmented terrestrial and shallow-marine environments. Furthermore, genetic and ecological studies increasingly suggest that species diversity is considerably higher in the pelagic realm than inferred from many morphological taxonomies.

Zoogeographic evidence suggests that ranges of many pelagic groups are much more limited by their ability to maintain viable populations than by any inability to disperse past tectonic and hydrographic barriers to population exchange. Freely dispersing pelagic taxa resemble airborne spores or wind-dispersed seeds that can drift almost anywhere but complete the entire life cycle only in favorable habitats. It seems likely that vicariant and allopatric models for speciation are far less important in pelagic evolution than sympatric or parapatric speciation in which dispersal is not limiting. Nevertheless, speciation can be quite rapid and involve cladogenesis even in cases where morphological data suggest gradual species transitions. Indeed, recent paleoecological and molecular studies increasingly suggest that classic examples of “phyletic gradualism” involve multiple, cryptic speciation events.

Paleoceanographic and climatic change seem to influence rates of turnover by modifying surface water masses and environmental gradients between them to create new habitats rather than by preventing dispersal. Changes in the vertical structure and seasonality of water masses may be particularly important since these can lead to changes in the depth and timing of reproduction. Long-distance dispersal may actually promote evolution by regularly carrying variants of a species across major oceanic fronts and exposing them to very different selection pressures than occur in their home range. High dispersal in pelagic taxa also implies that extinction should be difficult to achieve except though global perturbations that prevent populations from reestablishing themselves following local extinction. High rates of extinction in some pelagic groups suggests either that global perturbations are common, or that the species are much more narrowly adapted than we would infer from current taxonomies.

In this paper, I survey hindlimb and pelvic anatomy across non-avian dinosaurs and analyze these within a cladistic framework to quantify patterns of change within the locomotor apparatus. Specifically, I attempt to identify where homoplasy constitutes parallelism and may thereby be used to infer similar selective pressures on hindlimb function. Traditional methods of discrete character optimization are used along with two methods for evaluating changes in continuous characters in a phylogenetic context (squared-change parsimony and clade rank correlation). Resultant patterns are evaluated in light of the biomechanics of locomotion and the relationship between form and function in extant terrestrial vertebrates.

Although non-avian dinosaurian locomotor morphology is strikingly uniform, these analyses reveal the repeated derivations of several morphological features that have potential relevance for hindlimb locomotor function. Anterior and posterior iliac expansion, a medially oriented femoral head, and an elevated femoral lesser trochanter each evolved independently multiple times within Dinosauria. These changes probably reflect enlargement of several hindlimb muscles as well as a general switch in their predominant function from abduction-adduction (characteristic of “sprawling” limb postures) to protraction-retraction (characteristic of parasagittal, or “erect,” limb postures). Several “avian” characteristics are shared with more basal theropods, and many were acquired convergently in other dinosaurian lineages. The evolution of the avian hindlimb therefore represents a cumulative acquisition of characters, many of which were quite far removed in time and function from the origin of flight.

Living crocodilians (Crocodylia) and birds (Neornithes) differ in many aspects of hindlimb anatomy and locomotor function. How did this disparity evolve? We integrate information from fossils with functional descriptions of locomotion in living crocodilians and birds, using a phylogenetic perspective. We then outline the major changes in three-dimensional control of the hip joint along the line from the ancestral archosaur to birds. Our analysis reveals that most aspects of hip morphology and function in Alligator are ancestral for Archosauria. Femoral protractors and retractors are located cranial and caudal to the hip, respectively. Similarly, femoral adductors and abductors are located ventral and dorsal to the hip. Transformations of this ancestral pattern on the line to birds involved modifications in osteology, myology, and neural control. In some cases, homologous muscles changed function by acquiring new activity patterns. In others, activity was conserved, but origins and/or insertions were altered. Fossil theropods document the stepwise evolution of a novel mechanism of limb adduction/abduction involving long-axis rotation of the femur. This mechanism accounts for the conspicuous absence of significant musculature ventral and dorsal to the hip joint in extant birds.

Morphologically complex metazoans appear abruptly during the Cambrian explosion. Suggested measures of metazoan complexity include number of cell morphotypes and aspects of the genome such as the amount of DNA, the number of genes, and the information content of the genome or egg. Estimates of gene numbers are now available for metazoan species belonging to five different phyla or subphyla. There is little correlation between gene number and morphological complexity in the invertebrates: relatively complex forms can have fewer genes than relatively simple forms. Presumably, the more complex forms use more gene-expression events during development, implying that, on average, cis-regulatory elements of more complex invertebrates are richer in binding sites than are those of simpler forms. Vertebrates have many more genes than invertebrates and therefore have more total gene-expression events during development, although they may have, on average, fewer expression events per gene than the invertebrates. There are thus two genomic pathways in the evolution of metazoan complexity: one involves increasing the number of genes, the other involves increasing the number of cis-regulatory binding sites. Both modes were associated with the origin of bodyplans that first appear as fossils during the Cambrian explosion.

We compare refined data sets for Atlantic benthic foraminiferal oxygen isotope ratios and for North American mammalian diversity, faunal turnover, and body mass distributions. Each data set spans the late Paleocene through Pleistocene and has temporal resolution of 1.0 m.y.; the mammal data are restricted to western North America. We use the isotope data to compute five separate time series: oxygen isotope ratios at the midpoint of each 1.0-m.y. bin; changes in these ratios across bins; absolute values of these changes (= isotopic volatility); standard deviations of multiple isotope measurements within each bin; and standard deviations that have been detrended and corrected for serial correlation. For the mammals, we compute 12 different variables: standing diversity at the start of each bin; per-lineage origination and extinction rates; total turnover; net diversification; the absolute value of net diversification (= diversification volatility); change in proportional representation of major orders, as measured by a simple index and by a G-statistic; and the mean, standard deviation, skewness, and kurtosis of body mass. Simple and liberal statistical analyses fail to show any consistent relationship between any two isotope and mammalian time series, other than some unavoidable correlations between a few untransformed, highly autocorrelated time series like the raw isotope and mean body mass curves. Standard methods of detrending and differencing remove these correlations. Some of the major climate shifts indicated by oxygen isotope records do correspond to major ecological and evolutionary transitions in the mammalian biota, but the nature of these correspondences is unpredictable, and several other such transitions occur at times of relatively little global climate change. We conclude that given currently available climate records, we cannot show that the impact of climate change on the broad patterns of mammalian evolution involves linear forcings; instead, we see only the relatively unpredictable effects of a few major events. Over the scale of the whole Cenozoic, intrinsic, biotic factors like logistic diversity dynamics and within-lineage evolutionary trends seem to be far more important.

The morphological diversity of life has captivated systematists in the construction of classifications, embryologists in the study of development, and evolutionists in the formulation of theories of organic change. In a century marked by the advances of molecular biology, has the discipline of morphology produced anything … new? Yes. The solidification of paleontology and systematics and the emergence of macroevolution as a legitimate field owe much to an increased rigor in the analysis of morphological data. At the same time, the discipline of morphology has achieved an unprecedented sophistication through another development, the very expression of its maturity: theoretical morphology. Theoretical morphology forms the subject of McGhee's landmark book, an elegant combination of compendium and manifesto. Its richness and scope provide an opportunity for a critical appraisal of the discipline of morphology, particularly quantitative and developmental morphology.