Cheilostome bryozoans that grew as rigidly erect arborescent colonies dominate many bryozoan-rich assemblages of Tertiary age, in which they are found most commonly as small dissociated fragments. The regularity with which branching and branch thickening occur in intact colonies of living species provides a basis for quantitative reconstruction of these growth processes in fossils. We propose models to describe branch thickening, develop methods to extend both thickening and branching models to fossils, investigate the thickening and branching properties of four Paleocene and five Oligocene species and compare the properties of these fossils to those of nine living species.

The properties investigated are largely mutually independent and species specific irrespective of geologic age and have similar numerical ranges among different assemblages of coeval species. Species are evenly distributed across the range of possible morphologies between observed extremes, without obvious gaps. Statistically significant trends through time are identified in gradients of branch thickening, which have implications for the resistance of colonies to mechanical stress, and in angles of bifurcation, that are important in the way growing colonies occupy space.

In this paper we present a method for estimating soil pCO2 in ancient environments using the measured carbon-isotope values of pedogenic carbonates and plant-derived organic matter. The validity of soil pCO2 estimates proves to be highly dependent on the organic δ13C values used in the calculations. Organic matter should be sourced from the same paleosol profiles as sampled carbonates to yield the most reliable estimates of soil pCO2. In order to demonstrate the potential use of soil pCO2 estimates in paleoecological and paleoenvironmental studies, we compare samples from three Upper Jurassic localities. Soil pCO2 estimates, interpreted as a qualitative indicator of primary paleoproductivity, are used to rank the Late Jurassic terrestrial environments represented by the Morrison Formation in western North America, the informally named Lourinhã formation in Western Europe, and the Stanleyville Group in Central Africa. Because modern terrestrial environments show a positive correlation between primary productivity and faunal richness, a similar relationship is expected in ancient ecosystems. When the relative paleoproductivity levels inferred for each study area are compared with estimates of dinosaur generic richness, a positive correlation emerges. Both the Morrison and Lourinhã formations have high inferred productivity levels and high estimated faunal richness. In contrast, the Stanleyville Group appears to have had low primary productivity and low faunal richness. Paleoclimatic data available for each study area indicate that both productivity and faunal richness are positively linked to water availability, as observed in modern terrestrial ecosystems.

One of the most enduring evolutionary metaphors is Van Valen's (1973) Red Queen. According to this metaphor, as one species in a community adapts by becoming better able to acquire and defend resources, species with which it interacts are adversely affected. If those other species do not continuously adapt to compensate for this biotically caused deterioration, they will be driven to extinction. Continuous adaptation of all species in a community prevents any single species from gaining a long-term advantage; this amounts to the Red Queen running in place. We have critically examined the assumptions on which the Red Queen metaphor was founded. We argue that the Red Queen embodies three demonstrably false assumptions: (1) evolutionary adaptation is continuous; (2) organisms are important agents of extinction; and (3) evolution is a zero-sum process in which living things divide up an unchanging quantity of resources. Changes in the selective regime need not always elicit adaptation, because most organisms function adequately under many “suboptimal” conditions and often compensate by demonstrating adaptive flexibility. Likewise, ecosystems are organized in such a way that they tend to be robust and capable of absorbing invasions and extinctions, at least up to a point. With a simple evolutionary game involving three species, we show that Red Queen dynamics (continuous adaptation by all interacting species) apply in only a very small minority of possible outcomes. Importantly, cooperation and facilitation among species enable competitors to increase ecosystem productivity and therefore to enlarge the pool and turnover of resources. The Red Queen reigns only under a few unusual circumstances.

The centric diatom Stephanodiscus yellowstonensis Theriot and Stoermer is endemic to Yellowstone Lake, where it can be an important component of the summer phytoplankton assemblage. Its close relative, Stephanodiscus niagarae Ehrenberg, is abundant in nearby lakes and regional reservoirs. We used the stratigraphic record of Yellowstone Lake to investigate the evolution of S. niagarae to S. yellowstonensis and to describe the limnologic and climatic conditions associated with its evolution. A dramatic morphological shift took place between about 13.7 and 10.0 Ka, but morphology remained stable from 10 Ka to the present. Coincident with morphological change in the S. niagarae/S. yellowstonensis complex were changes in the diatom species assemblage, biogenic silica concentrations, sediment lithology, and regional vegetation. These changes suggest an environment that experienced progressive warming following the retreat of continental glaciers. We could not identify a specific selective factor driving evolution. Nevertheless, nonrandom morphological evolution strongly associated with continuous environmental change suggests that directional selection is a reasonable hypothesis to account for evolution of S. yellowstonensis. Protists are presumed to evolve gradually after speciation events because of large population size, high dispersal capacity, and low reproductive barriers. However, published diatom examples and the evolution of S. yellowstonensis suggest that it is premature to generalize about rates of evolution in protists, or at least to include diatoms in this generalization.

Micropaleontology is a peculiar subject: it is not easily defined, it focuses on geologic problems, and it ignores fundamental paleontologic and evolutionary questions it could best attack. As a result of its historic development, micropaleontology is directed to the solution of stratigraphic, paleoceanographic and paleoclimatologic problems, but it has seldom addressed paleobiologic or evolutionary ideas. It is a tradition rather than a discipline. The term “micropaleontology” and all it signifies should be abandoned, for it obscures natural relationships, attracts people with geologic rather than biologic approaches, isolates its practitioners in a blanket of systematics, biostratigraphies, and terminologies, and, as a result, discourages outsiders with other viewpoints or contributions from utilizing its fine fossil record. The growth of the field has been exponential in people-power and literature but not in the development of fundamentally new ideas. Micropaleontology has therefore contributed little to recent paleobiologic or evolutionary hypotheses, in spite of the possession by the many organisms relegated to it of biological properties and fossil records which have much potential for the generation and testing of such hypotheses. Microfossil studies have served geology powerfully and they should continue in that role, but they should also be used to fulfill their promise in the interpretation of paleobiogeography, paleoecology, morphology, evolutionary processes and the origin of new groups and ground plans.

656 mandibles and isolated teeth of the bovid Pachytragus solignaci Robinson from two localities in the Miocene Beglia Formation, Gafsa, Tunisia, have been individually aged on the basis of ontogenetic development and state of dental wear. The sample probably represents normal but seasonally restricted mortality. Life table analysis indicates an annual mortality rate of 25–30%, increasing in old animals. Dental dimensions are affected by stabilizing selection which sets in as the tooth comes into function, earlier mortality being nonselective in that respect.

A modern Lesser Flamingo (Phoeniconaias minor) assemblage was collected along the shoreline of Lake Emakat, a saline-alkaline lake in northern Tanzania. Taphonomic analysis found the assemblage to be heavily weathered. This is likely due to the bone's heightened exposure to solar radiation and corrosive soil and water chemistries, as is expected to occur in such depositional environments.

Analysis found that deep, wide, longitudinal cracks penetrate the medullar cavities of both weathered and unweathered long bones. The cause and taphonomic consequence of these cracks are addressed here, using data from Lake Emakat and from controlled studies. Results support repeated (episodic) submersion, followed by drying, as the causal mechanism behind these wet-dry cracks. Mineral salt uptake by bone may explain the early appearance and prevalence of these cracks in saline-alkaline lake settings, as compared to other depositional settings.

The rate of weathering and incidence of wet-dry cracking varies significantly across limb elements. This difference correlates to element specific resistance properties to external loading forces. Heavy weathering weakens the structural integrity of bone and can accelerate its fragmentation. This can lead to bird bone loss in nearshore and ephemeral wetland settings, which may then affect resulting skeletal part, diversity, and richness profiles. Heavy weathering can therefore obscure important taphonomic and paleoecological information.

The weathering data collected here are then applied to a fossil bird assemblage from the FLK Complex, (late Pliocene), Olduvai Gorge, in Tanzania. Results provide evidence for the effect of weathering on paleoecological and behavioral interpretations. Weathering should be considered when analyzing fossil bird assemblages.

We explore the relationships among the geographic ranges of genera, the ranges and positions of their constituent species, and the number of species they contain, considering variation among coeval genera and changes within genera over time. Measuring range size as the maximal distance, or extent, between occurrences within a taxon, we find that the range of the most widespread species is a good predictor of the range of the genus, and that the number of species is a better predictor still. This analysis is complicated by a forced correlation: the range of a genus must be at least as large as that of each of its constituent species. We therefore focus on a second measure of range, the mean squared distance, or dispersion, of occurrences from the geographic centroid, which, by analogy to the analysis of variance, allows the total dispersion of a genus to be compared to the mean within-species dispersion and the dispersion among species centroids. We find that among-species dispersion is the principal determinant of genus dispersion. Within-species dispersion also plays a major role. The role of species richness is relatively small. Our results are not artifacts of temporal variation in the geographic breadth of sampled data. The relationship between changes in genus dispersion and changes in within- and among-species dispersion shows a symmetry, being similar in cases when the genus range is expanding and when it is contracting. We also show that genera with greater dispersion have greater extinction resistance, but that within- and among-species dispersion are not demonstrable predictors of survival once the dispersion of the genus is accounted for. Thus it is the range of the genus, rather than how it is attained, that is most relevant to its fate. Species richness is also a clear predictor of survival, beyond its effects on geographic range.

The evolutionary history of shell geometry of Early Jurassic ammonoids during the Pliensbachian–Toarcian second-order mass extinction is explored at both adult and ontogenetic levels. The ontogenetic approach builds on the concept of allometric space to get insights into the developmental aspects of morphological evolution. Investigation of the deployment of taxa in adult morphospace and allometric space allows the appraisal of the temporal evolution of morphological and allometric disparities. Curves of taxonomic diversity, adult morphological disparity, allometric disparity, and average adult size are contrasted. Results show that during the Pliensbachian–Toarcian interval, ammonoids underwent two successive and drastic declines in taxonomic diversity. Patterns of morphospace and allometric space occupancy suggest nonselective extinction at both morphological and developmental levels. Another measure of allometric disparity suggests the occurrence of two heterochronic trends, a peramorphocline followed by a paedomorphocline, during the Toarcian. These trends are concomitant with changes in average adult size that compensate for the heterochronic effects and explain the striking stability of morphological disparity despite changes in diversity. The results also emphasize the existence of two contrasted evolutionary dynamics in Pliensbachian and Toarcian ammonoids. Methodologically, the allometric disparity approach appears to be a fruitful tool to analyze the rather understudied clade-wide ontogenetic aspects of morphological evolution. Combining multiple approaches to describe clade morphological dynamics leads to a better characterization and understanding of the diversity-disparities relationships and a better distinction of the potential processes driving these macroevolutionary patterns.

Here I test the hypothesis that temporal variation in geographic range size within genera is affected by the expansion and contraction of their preferred environments. Using occurrence data from the Paleobiology Database, I identify genera that have a significant affinity for carbonate or terrigenous clastic depositional environments that transcends the Database's representation of these environments during the stratigraphic range of each genus. These affinity assignments are not a matter of arbitrarily subdividing a continuum in preference; rather, genera form distinct, nonrandom subsets with respect to environmental preference. I tabulate the stage-by-stage transitions in range size within individual genera and the stage-by-stage changes in the extent of each environment. Comparing the two shows that genera with a preference for a given environment are more likely to increase in geographic range, and to show a larger average increase in range, when that environment increases in areal extent, and likewise for decreases in geographic range and environmental area. Similar results obtain for genera with preferences for reefal and non-reef settings. Simulations and subsampling experiments suggest that these results are not artifacts of methodology or sampling bias. Nor are they confined to particular higher taxa. Genera with roughly equal preference for carbonates and clastics do not have substantially broader geographic ranges than those with a distinct affinity, suggesting that, at this scale of analysis, spatial extent of preferred environment outweighs breadth of environmental preference in governing geographic range. These results pertain to changes over actual geologic time within individual genera, not overall average ranges. Recent work has documented a regular expansion and contraction when absolute time is ignored and genera are superimposed to form a composite average. Environmental preference may contribute to this pattern, but its role appears to be minor, limited mainly to the initial expansion and final contraction of relatively short-lived genera.

Kelps and other fleshy macroalgae—dominant reef-inhabiting organisms in cool seas—may have radiated extensively following late Cenozoic polar cooling, thus triggering a chain of evolutionary change in the trophic ecology of nearshore temperate ecosystems. We explore this hypothesis through an analysis of body size in the abalones (Gastropoda; Haliotidae), a widely distributed group in modern oceans that displays a broad range of body sizes and contains fossil representatives from the late Cretaceous (60–75 Ma). Geographic analysis of maximum shell length in living abalones showed that small-bodied species, while most common in the Tropics, have a cosmopolitan distribution, whereas large-bodied species occur exclusively in cold-water ecosystems dominated by kelps and other macroalgae. The phylogeography of body size evolution in extant abalones was assessed by constructing a molecular phylogeny in a mix of large and small species obtained from different regions of the world. This analysis demonstrates that small body size is the plesiomorphic state and largeness has likely arisen at least twice. Finally, we compiled data on shell length from the fossil record to determine how (slowly or suddenly) and when large body size arose in the abalones. These data indicate that large body size appears suddenly at the Miocene/Pliocene boundary. Our findings support the view that fleshy-algal dominated ecosystems radiated rapidly in the coastal oceans with the onset of the most recent glacial age. We conclude with a discussion of the broader implications of this change.

The most abundant and diverse graptolite assemblages are found in offshore, deep-water black shales—the classical “graptolite fades” (deep-water or isograptid biofacies). The mean duration of Ordovician graptolite species confined to the deep-water facies (here referred to as “group 1” species) is 2.19 Myr, significantly shorter than the mean duration of species in the deep-water facies that are also known in sediments of the shallow-water shelf or platform (“group 2” species) −4.42 Myr, indicating a significantly higher extinction probability (p = <0.001). These figures are based on the precise age ranges of species derived from the time-calibrated composite sequence of 1446 Ordovician to early Devonian graptolites, built by the constrained optimization procedure (CONOP) from 256 measured sections worldwide, and exclude the effects of the Hirnantian mass extinction. The difference between groups cuts across families, morphological types, and pandemic/endemic distributions. An environmental influence is strongly suggested, and although both groups were planktonic, they were unlikely to have shared the same habitat in the water column. The new duration measurements therefore are interpreted as favoring a depth-stratification of graptolite habitats in the water column.

Bryozoans are colonial animals and this permits the partitioning of their morphologic variability into components of within colony (i.e. within a single genotype) and between colony (i.e. between genotype) variance. These data have been obtained for four species of the endemic deep-sea genus Euginoma for a series of characters. In 8 comparisons, one component of the total variance dominated at the 5% level. Population (between colony) variance contributed significantly to the total variance in 63% of the comparisons (5 of 8); individual (within colony) variance contributed significantly to the total variance in 37% of the comparisons (3 of 8).

Compared to shallow water species, the surprising feature of the deep-sea data is that the between colony component of variance is as high as it is. Possibly in the more stable, deep-sea environment, the genotypic contribution to the variance of each individual colony is expressed to a greater degree than in the more variable, shallow water regime. If so, then analyses of variability in colonial animals may be an independent means of ascertaining stability gradients in the fossil record.

The unique holotype of the Triassic reptile Sharovipteryx (initially Podopteryx) mirabilis has been reexamined and redescribed, correcting the original account. It is a small lizard-like reptile with an elongate head and densely tubular femur and tibia each longer than the (estimated) intergirdle distance. (Part of the pectoral region has been lost and the remnants remain encased in the matrix.) The matrix retains the impression of portions of the integument, including a patagium that reached from the hindlimbs to the base of the tail. Unlike the original account, the patagium did not extend to the pectoral limbs. Experiments with models indicate that Sharovipteryx could have maintained a shallow glide if the femora were held at a shallow angle to the vertebral column and the tibia and feet extended out at right angles to it, thus stretching the integument. The pectoral limbs (with or without a fringing membrane) might have produced a variable canard; alternatively vertical bending of the tail could have applied drag, each making the glide more stable.

The Scleractinia, which are one of the most important builders of modern reefs, have been considered to have first appeared in the Middle Triassic. Recently, Paleozoic scleractiniamorphs have been reported from both the Ordovician and the Permian, suggesting that the scleractinian-like body plan was already established in the Paleozoic. Those Paleozoic scleractiniamorphs are considered either unsuccessful skeletonized offshoots (extinct experiments) or Paleozoic progenitors of the post-Paleozoic Scleractinia. Permian scleractiniamorphs are characterized by “ancestral” features and have no specific morphologies that deny scleractinian affinities. Molecular phylogenetics also indicate that extant scleractinians are monophyletic and originated long before their Triassic appearance. A Paleozoic origin for the Scleractinia is supported by morphological and molecular phylogenetic data. On the other hand, there is no positive evidence to show that different groups of scleractinians had separate soft-bodied precursors.

The Paleozoic scleractinians evolved within the framework of their basic body plan, and a direct derivation of the Scleractinia from the Rugosa is not probable. The Anthozoa are characterized by a bilaterally symmetrical body plan, which is traditionally considered to have been derived from other radially symmetrical Cnidaria. The problem of the origin of scleractinian body plan may provide a key for deciphering the early anthozoan radiation within the Bilateria. Other examples of Paleozoic Scleractinia and scleractiniamorphs will be found, probably in shallow-water reefal facies or deeper-water communities, bridging the stratigraphic gaps in occurrence and elucidating the origin of the Scleractinia and their body plan.

The stratigraphic distribution of graptoloid species within the upper Middle Ordovician strata of New York State represents a complex pattern of origination, extinction, faunal migration, and fluctuating relative abundances. In particular, the observed patterns of species turnover at graptolite biozone boundaries are apparently strongly correlated with lithofacies, sampling strategies, and the depositional effects of relative sea-level change.

Vertical facies changes that occur within third-order depositional sequences and fourth- or fifth-order parasequences are mirrored by changes in the graptoloid faunal composition. Large-scale faunal turnovers at biozone boundaries tend to occur either at sequence boundaries or at flooding surfaces within sequences (e.g., the base of Highstand System Tracts). The base of the Corynoides americanus and Climacograptus (D.) spiniferus Zones coincide with major onlap events, and the Orthograptus ruedemanni Zone fauna appears just below a Lowstand Systems Tract. Within individual biozones, smaller-scale changes such as the fluctuating relative abundances of graptoloid species coincide with higher-order parasequence cyclicity. These distribution patterns resemble recent computer-generated models for the sequence stratigraphic distribution of hypothetical taxa.

By combining good biogeographic control with a detailed sampling program, we are able to see through the patterns attributable to depositional cyclicity and identify the different components of faunal turnover (migration, speciation, and extinction).

Evolutionary trends observed over large clades have the potential to mask underlying trends that occur within their constituent subclades. A recent study of encephalization in the Caniformia (Carnivora, Mammalia) found evidence for an abrupt increase in median log-encephalization quotients (logEQs), indicating higher brain volume relative to body mass, at the end-Miocene, but gradual increase in the variance of logEQs. In this study, new endocranial volume estimates for fossil taxa in the well-sampled caniform subclade Canidae are reported. Using the encephalization data for the Canidae, hypotheses of evolution in encephalization allometries were tested with respect to canid phylogeny. The Akaike Information Criterion and likelihood ratios recovered support for a preferred hypothesis of the evolution of canid encephalization, which proposed two distinct allometric relationships: (1) a plesiomorphic grade of encephalization in the subfamilies Hesperocyoninae and Borophaginae and the paraphyletic canine genus Leptocyon, and (2) an apomorphic grade in the crown radiation of Caninae. This defines a shift in to higher encephalization, but without an associated change in the variance around the allometry. Increased canid encephalization coincides with a reorganization of the brain and the observed trend may reflect the evolution of complex social behavior in this clade.

Primary vascular architecture of members of the Paleozoic Aneurophytales (Progymnosper-mopsida) is described. This architecture is somewhat more complex but fundamentally similar (homologous) to that of members of the Trimerophytina, putative ancestors of aneurophytes. It is suggested that the presence of complex stelar morphology in aneurophytes was epiphenomenal, a passive result of changes in growth and development in a trimerophyte-like ancestor. Specifically, I suggest that the evolutionary transformation in primary vascular architecture from haplostele to ribbed protostele was a direct consequence of changes that affected the vertical spacing and degree of organization of lateral appendages in early vascular plants. This view is in sharp contrast to adaptationist explanations of change in stelar morphology expressed by other authors and provides an example of non-adaptive change in evolution.

In this paper we analyze the ontogenies of four species of pachypleurosaur (Reptilia: Sauropterygia) occurring in Triassic-age deposits in the Monte San Giorgio region, Switzerland. Preservation of multiple complete specimens representing a growth series from each taxon allows the comparison of ontogenetic trajectories through a space composed of nine variables important in the evolution of the clade. Trajectories are characterized using the multivariate generalization of the allometry equation and then compared through calculations of the angles between allometry vectors. Individual coefficients of vectors are compared after calculation of bootstrapped confidence intervals. Pachypleurosaur ontogeny is found to be allometric and generally conserved, although significant differences between taxa exist. Shape-disassociated allometric changes characterize the transition between Serpianosaurus and Neusticosaurus, while allometric changes within Neusticosaurus are less significant. N. edwardsii is inferred to have arisen through hypermorphosis. Interpretation of whole-body heterochrony in multivariate analysis is discussed.