Digitizing the third upper molars of Microtus pennsylvanicus reveals evolutionary change in some traits but stability in others during the last 30,000 years. Fossils from Strait Canyon, Virginia (radiocarbon dated between 29,870 + 1,800/-1,400 and 17,880 ± 150 yr B.P.) were compared with modern samples from the margin of the species' conterminous geographic range: Virginia, Colorado, Alaska, and Hudson Bay. Since the late Pleistocene a modification from narrow to wide teeth took place in the eastern populations but not in the western ones, and populations in Virginia apparently evolved wider confluence between triangles 1 and 2. Traits that remained stable through time in the Virginia area, but not necessarily elsewhere, include the numerical shape factors of the occlusal surface and the posterior loop. Potentially widespread stability is evident in the perimeter of the posterior loop relative to total tooth perimeter. Tooth length forms a geographic cline, in which both the fossil and modern populations from Virginia occupy the same position. A peripheral population from southern Colorado shows the most derived dental morphology. These morphological relationships suggest that mosaic evolution operates in arvicoline lineages and provide testable paradigms for future studies.

Although morphological variation is known to influence the evolutionary fates of species, the relationship between morphological variation and survivorship in the face of extinction-inducing perturbations is poorly understood. Here, we investigate this relationship for veneroid bivalves in association with the Plio-Pleistocene extinction in Florida. Fourteen pairs of related species were selected for analysis, with each pair including one species that survived the Plio-Pleistocene extinction and another that became extinct during the interval. Morphological landmark data were acquired for more than 1500 museum specimens, representing 19 localities that encompass four well-known Plio-Pleistocene units in the study region. Procrustes superimposition was applied to each sample, and overall multivariate variation was calculated as the mean squared partial Procrustes distance between specimens and their mean form. Morphological variation was calculated at three geographic scales for each species, and differences in variation between survivors and victims were examined within each species pair. Results indicate that species surviving the Plio-Pleistocene extinction were significantly more variable morphologically than victims. Greater morphological variation may promote survivorship by directly enhancing species adaptations to changing conditions or by permitting the occupation of a larger geographic range. Alternatively, high morphological variation and survivorship may both be mediated by a third variable, such as large geographic range.

We use Fourier analysis to investigate evolutionary dynamics related to periodicities in marine fossil biodiversity. Coherent periodic fluctuation in both origination and extinction of “short-lived” genera (those that survive <45 Myr) is the source of an observed ∼62 million year periodicity (which we confirmed in an earlier paper and also found to be ubiquitous in global compilations of marine diversity). We also show that the evolutionary dynamics of “long-lived” genera (those that survive >45 Myr) do not participate in the periodic fluctuation in diversity and differ from those of “short-lived” genera. The difference between the evolutionary dynamics of long- and short-lived genera indicates that the periodic pattern is not simply an artifact of variation in quality of the geologic record. Also, the interplay of these two previously undifferentiated systems, together with the secular increase in abundance of “long-lived” genera, is probably the source of observed but heretofore unexplained differences in evolutionary dynamics between the Paleozoic and post-Paleozoic as reported by others. Testing for cycles similar to the 62-Myr cycle in fossil biodiversity superimposed on the long-term trends of the Phanerozoic as described in Paper I, we find a significant (but weaker) signal in sedimentary rock packages, particularly carbonates, which suggests a connection. The presence of a periodic pattern in evolutionary dynamics of the more vulnerable “short-lived” component of the marine fauna demonstrates that a long-term periodic fluctuation in environmental conditions capable of affecting evolution in the marine realm characterizes our planet's history. Coincidence in timing is more consistent with a common cause than with sampling bias. A previously identified set of mass extinctions preferentially occur during the declining phase of the 62-Myr periodicity, supporting the idea that the periodicity relates to variation in biotically important stresses. Further work should focus on finding links to physical phenomena that might reveal the causal system or systems.

Oysters, whose inner shell layer contains chambers, vesicles, and sometimes chalky deposits, often have extraordinarily thick shells of large size, prompting the idea that there is something unusual about the process of shell fPormation in these and similarly structured bivalves with the oyster syndrome. I propose the hypothesis that calcifying microbes, especially sulfate-reducing bacteria growing on organic substrates in fluid-filled shell-wall chambers, are responsible for shell calcification away from the shell-secreting mantle of the host bivalve. Other phenomena, including the formation of cameral deposits in fossil cephalopods, the cementation of molluscs and barnacles to hard substrata, the formation of a calcified intriticalx on the shell's exterior, and cementation of objects by gastropods on the shell for camouflage, may also involve calcifying bacteria. Several lines of inquiry are suggested to test these hypotheses.

Bases of attachment of Archimedes colonies are very rare, but many colonies show evidence of origination from a pre-existing set of branches. The evidence consists of spiralled supports (axial screws) originating from fragments of pre-existing sets of branches, proximal taper of colonies or axial screws to a diameter smaller than that of bases of attachment, early stages of new colonies originating from the whorl margins of older colonies, and sets of laterally fused colonies. In the most favorable environments Archimedes populations were dominated by a few genets that consisted of hundreds to perhaps thousands of genetically identical colonies.

The ability to reproduce by colony fragmentation was probably an important cause of Archimedes' abundance in the Chesterian seas of eastern North America, especially where it occurred in dense populations in the lee of submarine carbonate shoals. Colony fragmentation, the importance of which is becoming increasingly recognized in modern corals and bryozoans, was also of importance in Paleozoic seas.

It has recently been argued that barren intervals of marine sedimentary rock are less common in the Cenozoic than in the Paleozoic, and that this arises as a direct consequence of widespread epeiric seas and the prevalence of dysaerobic conditions at such times. We show, using an independent and more direct measure of rock outcrop through time in western Europe, that barren marine sedimentary rocks do become less frequent toward the present, but that this is not linked to any epeiric-seas effect. The proportion of barren to fossiliferous rock outcrop correlates well with the inferred Phanerozoic marine diversity curve (although more so in the Paleozoic than in the post-Paleozoic), and shows no correlation or only a weak negative correlation with area over which the sediments have been deposited. We therefore concluded that the Phanerozoic trend in fossiliferousness most likely records the degree to which space is occupied in the shallow marine realm.

Parasitic trematode worms leave characteristic pits in their bivalve mollusk hosts and represent an ideal system for analyzing parasite-host interactions through space and time with statistically meaningful sample sizes. Previous work in Late Pleistocene–Holocene sequences from the Po plain revealed significant long-term fluctuations in trematode prevalence values: higher prevalence in retrogradational environments (TST) and negligible prevalence in progradational environments (HST). Here we expand upon this work by investigating traces of parasitism, kleptoparasitism, and predation on mollusk death assemblages from two domains along the northern Adriatic coastline. The domain north of the Po delta (TST-like) and the southern domain (including the Po delta; HST-like) comprise environments comparable to those recovered in late Holocene (<6 Kyr) subsurface progradational deposits. We collected 17,299 specimens representing 111 species from 11 locations on the northern Adriatic coast of Italy. Our results reveal high predation pressure, a high diversity of host taxa, and widespread presence of trematode infestation in starved, oligotrophic, environmentally more stable (i.e., TST-like) settings north of the Po delta. Immediately south of the Po delta, in settings with strong and variable sedimentary input, almost no infestation is recorded. The reappearance of infestation is evident in the southern portion of the study area (i.e., Cattolica-Montemarciano), relatively far from the highly stressed environments south of the Po River. There is no significant difference in trematode prevalence values between fossil and modern samples. The distribution of spionid traces (an indicator of stressed environments) was nearly the opposite of that displayed by trematodes. Drilling frequency is highest in TST-like environments and is not correlated with diversity indices. These results suggest that temporal trends of trematode prevalence (and possibly also other biotic interactions) in sedimentary successions are controlled by environmental changes driven by glacio-eustatic dynamics, and reaffirm the importance of interpreting temporal trends in the context of spatial variation.

Issues of directionality in the history of life can be framed in terms of six major evolutionary steps, or megatrajectories (cf. Maynard Smith and Szathmáry 1995): (1) evolution from the origin of life to the last common ancestor of extant organisms, (2) the metabolic diversification of bacteria and archaea, (3) evolution of eukaryotic cells, (4) multicellularity, (5) the invasion of the land and (6) technological intelligence. Within each megatrajectory, overall diversification conforms to a pattern of increasing variance bounded by a right wall as well as one on the left. However, the expanding envelope of forms and physiologies also reflects—at least in part—directional evolution within clades. Each megatrajectory has introduced fundamentally new evolutionary entities that garner resources in new ways, resulting in an unambiguously directional pattern of increasing ecological complexity marked by expanding ecospace utilization. The sequential addition of megatrajectories adheres to logical rules of ecosystem function, providing a blueprint for evolution that may have been followed to varying degrees wherever life has arisen.

The placement of angiosperms and Gnetales in seed plant phylogeny remains one of the most enigmatic problems in plant evolution, with morphological analyses (which have usually included fossils) and molecular analyses pointing to very distinct topologies. Almost all morphology-based phylogenies group angiosperms with Gnetales and certain extinct seed plant lineages, while most molecular phylogenies link Gnetales with conifers. In this study, we investigate the phylogenetic signal present in published seed plant morphological data sets. We use parsimony, Bayesian inference, and maximum-likelihood approaches, combined with a number of experiments with the data, to address the morphological–molecular conflict. First, we ask whether the lack of association of Gnetales with conifers in morphological analyses is due to an absence of signal or to the presence of competing signals, and second, we compare the performance of parsimony and model-based approaches with morphological data sets. Our results imply that the grouping of Gnetales and angiosperms is largely the result of long-branch attraction (LBA), consistent across a range of methodological approaches. Thus, there is a signal for the grouping of Gnetales with conifers in morphological matrices, but it was swamped by convergence between angiosperms and Gnetales, both situated on long branches. However, this effect becomes weaker in more recent analyses, as a result of addition and critical reassessment of characters. Even when a clade including angiosperms and Gnetales is still weakly supported by parsimony, model-based approaches favor a clade of Gnetales and conifers, presumably because they are more resistant to LBA. Inclusion of fossil taxa weakens rather than strengthens support for a relationship of angiosperms and Gnetales. Our analyses finally reconcile morphology with molecules in favoring a relationship of Gnetales to conifers, and show that morphology may therefore be useful in reconstructing other aspects of the phylogenetic history of the seed plants.

A number of natural events can cause ozone depletion, including asteroid and comet impacts, large-scale volcanism involving the stratospheric injection of chlorine, and close cosmic events such as supernovae. These events have previously been postulated to have been sole or contributory causes of mass extinctions. Following such events, UV-B radiation would have been elevated at the surface of the earth. The possibilities for detecting elevated UV-B as a kill mechanism in the fossil record are discussed. In the case of impact events and large-scale volcanism, the taxa affected by increases in UV-B radiation are likely to be similar to those affected by cooling and by the initial drop in irradiance caused by stratospheric dust injection. Thus UV-B may synergistically exacerbate the effects of these other environmental changes and contribute to stress in the biosphere, although UV-B alone is unlikely to cause a mass extinction. By the same token, however, this similarity in affected taxa is likely to make delineating the involvement of UV-B radiation in the fossil record more difficult. Cosmic events such as supernovae may produce smaller extinction events, but ones that are “cleaner” UV catastrophes without the involvement of other environmental changes.

Organismal metabolic rates reflect the interaction of environmental and physiological factors. Thus, calcifying organisms that record growth history can provide insight into both the ancient environments in which they lived and their own physiology and life history. However, interpreting them requires understanding which environmental factors have the greatest influence on growth rate and the extent to which evolutionary history constrains growth rates across lineages. We integrated satellite measurements of sea-surface temperature and chlorophyll-a concentration with a database of growth coefficients, body sizes, and life spans for 692 populations of living marine bivalves in 195 species, set within the context of a new maximum-likelihood phylogeny of bivalves. We find that environmental predictors overall explain only a small proportion of variation in growth coefficient across all species; temperature is a better predictor of growth coefficient than food supply, and growth coefficient is somewhat more variable at higher summer temperatures. Growth coefficients exhibit moderate phylogenetic signal, and taxonomic membership is a stronger predictor of growth coefficient than any environmental predictor, but phylogenetic inertia cannot fully explain the disjunction between our findings and the extensive body of work demonstrating strong environmental control on growth rates within taxa. Accounting for evolutionary history is critical when considering shells as historical archives. The weak relationship between variation in food supply and variation in growth coefficient in our data set is inconsistent with the hypothesis that the increase in mean body size through the Phanerozoic was driven by increasing productivity enabling faster growth rates.

Decay experiments are becoming a more widespread tool in evaluating the fidelity of the fossil record. Character interpretations of fossil specimens stand to benefit from an understanding of how decay can result in changes in morphology and, potentially, total character loss. We performed a decay experiment for the Class Enteropneusta to test the validity of anatomical interpretations of the Burgess Shale enteropneust Spartobranchus tenuis and to determine how the preservation of morphological features compares with the sequence of character decay in extant analogues. We used three species of enteropneust (Saccoglossus pusillus, Harrimania planktophilus, and Balanoglossus occidentalis) representing the two major families of Enteropneusta. Comparisons between decay sequences suggest that morphological characters decay in a consistent and predictable manner within Enteropneusta, and do not support the hypothesis of stemward slippage. The gill bars and nuchal skeleton were the most decay resistant, whereas the gill pores and pre-oral ciliary organ were unequivocally the most decay prone. Decay patterns support the identification of the nuchal skeleton, gill bars, esophageal organ, trunk, and proboscis in Spartobranchus tenuis and corroborate a harrimaniid affinity. Bias due to the taphonomic loss of taxonomically informative characters is unlikely. The morphologically simple harrimaniid body plan can be seen, therefore, to be plesiomorphic within the enteropneusts. Discrepancies between the sequence of decay in a laboratory setting and fossil preservation also exist. These discrepancies are highlighted not to discredit the use of modern decay studies but rather to underline their non-actualistic nature. Paleoenvironmental variables besides decay, such as the timeframe between death and early diagenesis as well as postmortem transport, are discussed relative to decay data. These experiments reinforce the strength of a comprehensive understanding of decay sequences as a benchmark against which to describe fossil taxa and understand the conditions leading to fossilization.

Spinal ossified tendons are a defining character for Ornithischia, one of the two major clades of dinosaurs. The function of these bony rods has remained a mystery since their first detailed description in 1886. Qualitative approaches to understand ossified tendon function have resulted in different ecological and behavioral interpretations for ornithopod dinosaurs. To evaluate ossified tendon function, this study constructed finite element models of the vertebral column for two ornithopod taxa: Tenontosaurus, which shows the plesiomorphic condition of longitudinally arrayed tendons along the spinous processes, and Brachylophosaurus, which exhibits a lattice of tendons along the spinous processes. Both models predict that ossified tendons stiffened the vertebral column, especially the tail, but the derived lattice of ossified tendons in iguanodontoidean dinosaurs, like Brachylophosaurus, increased spinal stiffness more than the plesiomorphic condition. Caudofemoral muscles that retracted the hindlimb during locomotion attached the femur to the tail in ornithopods. Increased tail stiffness caused by intratendinous ossification may have influenced locomotion by rigidly anchoring M. caudofemoralis longus to the tail, thereby allowing a more forceful retraction of the hindlimb by reducing ventral flexion of the tail during muscle contraction. Ossified tendons may also have been important for storing elastic energy throughout the gait cycle. Moreover, the lattice of ossified tendons stiffened the trunk and tail nearly equally in Brachylophosaurus, indicating the evolution of a postural function by passively supporting the epaxial musculature in maintaining a horizontal vertebral column.

The set of environmental conditions under which a taxon can survive and maintain viable populations, known as the ecological niche, is a fundamental determinant of a taxon's distribution. Because of the central importance of ecological niches, they have been assumed to remain relatively stable during intervals of morphological stasis. However, the assumption of niche stability has rarely been tested directly with fossil data spanning multiple temporal intervals. Thus, the conditions under which this assumption is likely to be accurate are not well understood. In this study, we use ecological niche modeling (ENM) to reconstruct the ecological niche for 11 genera of marine benthos (crinoids, trilobites, molluscs, bryozoans, and corals) from the Type Cincinnatian Series (Late Ordovician, Katian Stage) across nine temporal intervals spanning approximately three million years. This interval includes both abiotic environmental change (gradual sea-level fall) and biotic change (rapid pulses of the Richmondian Invasion), thus allowing the relative effect of different environmental perturbations to be constrained. A previous symmetrical analysis of niche stability of brachiopod species recovered an increase in niche evolution following the Richmondian Invasion. Herein we test the generality of the brachiopod pattern within the community. Niche stability was evaluated in geographic space, ecological space, and niche parameter space. Niche stability varied through time; during the Pre-Invasion interval, taxa exhibited niche stability during gradual shallowing of sea level in the basin, whereas niche evolution became more common during the Richmondian Invasion. Taxa adjusted to the increased competition by altering aspects of their niche. Notably, surviving taxa contracted their niche into a subset of their previous niche parameters. This represents an adaptive response to increased competition for resources with the newly established invader taxa, and it was employed most successfully by generalist taxa. Patterns of niche evolution were congruent between clades, among feeding styles, and across taxonomic levels.

Given estimates of the variation in total standing species richness through the periods of the Phanerozoic, mean species duration, and the relative intensity of the sampling of the fauna from each of the periods, the expected number of described species can be predicted for each period of the Phanerozoic using an analytic sampling model. This model is based on the assumption that the relative abundances of species in any geologic period can be approximated by the canonical (lognormal) species-abundance distribution.

Three commonly cited models of standing species richness (Valentine, 1973; Gould et al., 1977; Bambach, 1977) each suggest different patterns of species richness in the Phanerozoic. By assuming that sampling of the fossil record is proportionate to sediment volume, it can be shown with the sampling model that the Empirical, Equilibrium, and Species-Richness Models each predict that the number of described species will be strongly correlated with sediment volume. Equally high correlations are predicted if it is assumed that sampling is proportionate to sediment area or to paleontological interest. The correlations predicted for each of the three models are remarkably similar. The impact of sampling effects is so strong that the variations in species richness postulated by these three models are almost completely obscured. Preservational biases will probably only further obscure the relationship between the number of described species and total species richness. Therefore, it seems likely that analysis of trends in the total number of described species will be of little use in determining trends in worldwide species richness in the Phanerozoic.

Comparison of the actual patterns of variation in the number of described species and the expected numbers of described species predicted by the sampling model reveals that more species are known from the Cenozoic than would be predicted from the abundance of Cenozoic sediments or from the amount of paleontological interest in the Cenozoic. This might have resulted from the Cenozoic sediments remaining relatively free of diagenetic effects which might have destroyed the fossils entombed in the sediments.

The hypothesis is proposed that mean level of heterozygosity is functionally related to rate of speciation in evolutionary phylads. Under this hypothesis, phylads which speciate more rapidly do so because of increased level of within-species genetic variability which is then available to conversion to species differences under appropriate ecological or environmental conditions. An important corollary is that rate of speciation could be limited in phylads with low genetic variability, irrespective of environmental considerations.

This hypothesis has been tested with respect to electrophoretically detectable variation in products of structural genes in two families of North American fishes characterized by grossly different rates of speciation. Totals of 69 species of the highly speciose Cyprinidae, and 19 species of the relatively depauperate Centrarchidae, were assayed for mean level of heterozygosity at 11–24 genetic loci. Since Cyprinidae and Centrarchidae exhibit on the average nearly identical levels of genic variation (Ĥ = 0.052 ± 0.004, and Ĥ = 0.049 ± 0.009, respectively), the hypothesis that level of heterozygosity affects rate of speciation in these fishes is not supported.

Nonetheless, the amount of genic variability in both Cyprinidae and Centrarchidae is large, comparable to mean levels in previously studied vertebrates. The great wealth of genome variability, reflected in the electrophoretic variation present in virtually all outcrossing organisms, apparently can accommodate considerable flexibility in rate and pattern of evolutionary response to the various environmental regimes challenging organisms.

The exceptionally preserved Triassic (Anisian) Grès à Voltzia fauna from the Vosges of northeastern France is compared with four major Carboniferous Konservat-Lagerstätten. The problem of comparing faunas with different types of preservation and degrees of taxonomic determination is addressed with a newly devised similarity coefficient. This coefficient quantifies and combines data from different levels in the taxonomic hierarchy and allows comparisons between biotas in a consistent direction regardless of relative diversity. The rank order of similarity between the four Carboniferous Konservat-Lagerstätten and the Grès à Voltzia is as follows: 1. Mazon Creek (Westphalian D) of Illinois; 2. Glencartholm (Visean) of southern Scotland; 3. Bear Gulch (Namurian) of Montana; and 4. Blanzy-Montceau (Stephanian) of France. These occurrences represent conditions transitional between nearshore fully marine and fresh water. The Grès à Voltzia fauna is significantly closer to the Mazon Creek fauna than to the others; the taxonomic overlap with Blanzy-Montceau and Bear Gulch is limited.

Stratigraphic age has an insignificant influence on the result; indeed, the fauna closest in age to the Grès à Voltzia, that of Blanzy-Montceau, is least similar. Taphonomic factors are important in determining the range of organisms preserved. The Glencartholm fauna is represented only by forms with either mineralized or robust chitinous skeletons, implying a greater degree of decay prior to the onset of diagenetic mineralization than in the other Konservat-Lagerstätten. Environment, however, is the major control on similarity. The Mazon Creek biota, like that of Grès à Voltzia, represents settings transitional between terrestrial and marine-influenced delta. Groups common to both Konservat-Lagerstätten include medusae, brachiopods, polychaetes, bivalve and gastropod molluscs, limulids, scorpions, spiders, branchiopods, ostracodes, malacostracans, cycloids, euthycarcinoid and myriapod uniramians, insects, fish, and tetrapods. There is a striking continuity between the faunas of Carboniferous and Triassic transitional sedimentary environments. Groups that were adapted to fluctuating conditions (e.g., shifting salinity) show strong congruence at the family and lower levels and were little affected by Permian extinctions. The major taxonomic contrasts are in the eumalacostracans and insects: many of the groups represented in the Grès à Voltzia appeared in the Permian and radiated across the Permo-Triassic boundary as Paleozoic forms became extinct.

Truth, goes an old proverb, is the daughter of time. Fifty years ago, G. G. Simpson (1944) brought paleontology into the Neodarwinian fold, arguing that evolutionary tempo can be documented in the geological record and used to inform debate about evolutionary mode. Today, increasingly sophisticated paleontological investigations of rate—be it diversification, extinction, migration, morphological change, or divergence in macromolecular sequence—require calibration of the geological time scale with a precision far greater than Simpson could have anticipated. Expanding research on the relationships between environmental history and evolution also requires unprecedented resolution in correlation and geochronometry.

Several groups of siliceous microorganisms possess a skeletal latticework of interconnected rods. Skeletal configurations of one of these groups, the silicoflagellates, can be produced by a simple mathematical model that minimizes the apical surface area for a given basal area and internal volume. A similar model that minimizes the total length of the skeletal elements, and thus the silica utilization and skeletal weight, produces configurations that are generally less common in silicoflagellates. The diversity of silicoflagellate skeletal morphologies suggests that both the minimization of apical surface area and the conservation of skeletal material may be important factors in skeletal design. The two most important morphologies found in modern oceans, the four-sided Dictyocha and the six-sided Distephanus, can co-occur in an environment where both factors have some relative importance. However, these models do not explain the range of silicoflagellate skeletal morphology found in nature.

It has been suggested that a sudden event at the end of the Cretaceous period caused a major extinction that was felt disproportionately by creatures in the water column. It also has been argued that benthic deposit feeders, being relatively independent of abundance of organic particles in the water column, should have survived the crisis more readily than suspension feeders, which depended more upon feeding upon phytoplankton. I argue that the hypothesis of relative immunity of deposit feeders is insufficient, because deposit feeders by and large depend upon a supply of organic matter from the water column and would have succumbed to food shortage nearly as rapidly as suspension feeders, possibly within a maximum of three to six months. This near simultaneity of extinction would have been especially true of continental shelf environments. Even in some parts of the deep sea, it is likely that a dependence upon the water column above might have caused deep-sea deposit feeders to succumb rapidly. Therefore, deposit feeders would not necessarily have outsurvived suspension feeders during a crisis of depleted water-column phytoplankton, increased shading by inert particles, or poisoning of the water column and killing of phytoplankton. The relatively lower rate of extinction of nuculoid bivalves may relate instead to their presence in deeper-water refuge habitats, their apparent relative ability to diversify in higher latitudes, or their resistance to factors other than food shortage.