Long-term persistence of patterns in geographic variation within species is an interesting and puzzling phenomenon. I present a well-defined natural experiment in the land snail Cerion agassizi from the islands of Great Bahama Bank. C. agassizi is the best-known fossil of the ca. 120,000 years BP dunes of New Providence, Cat and Eleuthera Islands; populations have survived on Cat and Eleuthera. During the Wisconsin glacial advance, all these islands joined together in an emergent bank. Presence of the same species on two islands at two times permits a test for both time signatures (does change occur in the same manner on both islands) and island signatures (do aspects of shell phenotypes remain constant on each island through time).

Factor and discriminant analyses establish morphological separations among fossil populations of the three islands. These differences occur along pathways specified by well-known covariance sets in the complex allometric ontogeny of Cerion. By these routes, small variations in the geometry of growth may be magnified to large differences in external appearance. I found a time signature, probably attributable to introgression of modern populations by Cerion glans on both Cat and Eleuthera. Despite the intermediate period of emergence and joining of all islands, I also found an island signature in the preservation through time, on both Cat and Eleuthera, of the differentia that separate fossil populations. The basic distinctions of the two islands, expressed as patterns of covariance in growth, have been stable for at least 120,000 years.

The species richness of extant metazoan phyla displays a strong association with the ontogenetic timing of germ-line determination. Taxa with early ontogenetic determination of the germ-line are characterized by low species number relative to taxa with late or variable determination. A test for a similar association in fossil taxa is compromised by the fact that all phyla displaying early determination of the germ-line are small, soft-bodied groups which lack a reliable fossil record. To the extent that this correlation reflects causality, patterns of evolutionary diversification have been decidedly non-random. The fact that ontogenetic timing of germ-line determination defines the extent to which genetic variation arising during the course of ontogeny may be inherited suggests a possible causal foundation for the pattern.

Global taxonomic richness is affected by variation in three components: within-community, or alpha, diversity; between-community, or beta, diversity; and between-region, or gamma, diversity. A data set consisting of 505 faunal lists distributed among 40 stratigraphic intervals and six environmental zones was used to investigate how variation in alpha and beta diversity influenced global diversity through the Paleozoic, and especially during the Ordovician radiations. As first shown by Bambach (1977), alpha diversity increased by 50 to 70 percent in offshore marine environments during the Ordovician and then remained essentially constant for the remainder of the Paleozoic. The increase is insufficient, however, to account for the 300 percent rise observed in global generic diversity. It is shown that beta diversity among level, soft-bottom communities also increased significantly during the early Paleozoic. This change is related to enhanced habitat selection, and presumably increased overall specialization, among diversifying taxa during the Ordovician radiations. Combined with alpha diversity, the measured change in beta diversity still accounts for only about half of the increase in global diversity. Other sources of increase are probably not related to variation in gamma diversity but rather to appearance and/or expansion of organic reefs, hardground communities, bryozoan thickets, and crinoid gardens during the Ordovician.

Analysis of the stratigraphic records of 19,897 fossil genera indicates that most classes and orders show largely congruent rises and falls in extinction intensity throughout the Phanerozoic. Even an ecologically homogeneous sample of reef genera shows the same basic extinction profile. The most likely explanation for the congruence is that extinction is physically rather than biologically driven and that it is dominated by the effects of geographically widespread environmental perturbations influencing most habitats. Significant departures from the congruence are uncommon but important because they indicate physiological or habitat selectivity. The similarity of the extinction records of reef organisms and the marine biota as a whole confirms that reefs and other faunas are responding to the same history of environmental stress.

The species-level properties of geographic range and geologic duration are often used as variables in evolutionary studies. However, estimates of species duration are not independent of estimates of geographic range. Before these properties are used in macroevolutionary hypotheses, error associated with these estimates must be quantified. This error may lead to spurious inferences of evolutionary processes. To assess the error associated with estimates of geographic range and geologic duration, we modeled various sampling regimes and calculated the bias associated with these estimates.

We present three analyses which document the bias associated with estimates of geographic range and geologic duration. First, we find a positive correlation between local abundance and geographic range for a sample of 180 species of Recent prosobranch gastropods from the northeastern temperate Pacific Ocean. Therefore, geographically short-ranging species are less likely to be represented in the fossil record than geographically long-ranging species because of their local rarity. Second, we demonstrate that the chance of underestimating the geographic range of a species is acute for species with restricted spatial distributions, further compounding the problem of documenting their distribution in space and time. Third, we present a simulation which quantifies the degree of autocorrelation between geographic range and geologic duration for different levels of sampling resolution and spatial distributions of fossil localities.

We propose that the kelps (Laminariales) radiated in the North Pacific following the onset of late Cenozoic polar cooling. The evidence is that (1) extant kelps occur exclusively in cold-water habitats; (2) all but one of 27 kelp genera occur in the North Pacific, 19 of these exclusively; and (3) limpets and herbivorous marine mammals obligately associated with kelps or other stipitate brown algae appeared late in the Cenozoic, even though more generalized forms of both groups are much older. We propose, further, that sea otters and perhaps other groups of benthic-feeding predatory mammals, whose late Cenozoic distributions all were limited to the North Pacific, created an environment for the evolution of kelps in which the intensity of herbivory was unusually low. We hypothesize that this interaction created predictable differences among habitats in the intensity of herbivory on several spatial scales, with resulting trade-offs between anti-herbivore defenses and plant competitive abilities in their respective floras. Sea otters incur time and energy costs for diving, resulting in depth-related reductions to foraging efficiency and thus increased sizes and densities of herbivorous sea urchins. Thus, the deep-water flora is well defended, but competitively subordinate, compared with the shallow-water flora. Similarly, we argue that during the same period of earth history, predation had less of a limiting influence on herbivorous invertebrates in the temperate southwestern Pacific. We hypothesize that (1) consequent biogeographical differences in the intensity of herbivory may have selected the phenolic-rich brown algal flora in temperate Australia/New Zealand; and (2) tightly coevolved plant/herbivore interactions may explain why Australian and New Zealand herbivores are undeterred by phenolics and why other classes of secondary compounds in the Australian/New Zealand flora significantly deter herbivores.

Being of especially high quality, the Neogene fossil record of planktonic foraminifera offers special opportunities for assessing patterns of extinction and speciation. A variety of metrics indicates that within this group the mean duration of lineages has been much shorter (rate of extinction has been higher) for the globorotaliid clade than for the globigerinid clade. Furthermore, in the globorotaliid clade rates of extinction and speciation have not been closely linked to changes in diversity, but rather have been relatively high even at times when diversity has undergone little change. Thus, the globorotaliid clade has undergone more rapid evolutionary turnover than the globigerinid clade. Data for living species reveal that neither geographic range nor temperature tolerance is the primary factor controlling lineage duration. On the other hand, there is evidence that lineages marked by low abundance (small population size) are relatively short-lived. The reason that globorotaliid lineages have generally survived for shorter intervals, on the average, may be that their populations have been less abundant and less stable. Usually they live deeper in the water column, where food is often sparse, and many flourish only in areas of upwelling. Furthermore, the globorotaliids lack symbiotic algae for nutritional support. The same ecological factors may have accelerated speciation in the globorotaliid clade, by causing species to be patchily distributed. Thus, population size and structure have been more important than geographic range in determining rates of extinction and speciation. This parallels the situation for Neogene marine bivalves.

For planktonic foraminifera, as for Western Atlantic Bivalvia, the normal pattern of extinction was reversed in late Pliocene time, apparently in response to climatic cooling. The globigerinids suffered a sudden pulse of extinction. The deeper dwelling globorotaliids fared better; probably many of their species benefited from elevation of the seasonal thermocline into the photic zone. At the same time, rate of speciation declined in the globorotaliid clade, which supports the idea, inferred from the evolutionary history of marine bivalves, that an increase in the size and stability of populations should depress both rate of extinction and rate of speciation.

A method is provided to calculate the expected distribution of species' ranges for use as a basis of comparison for paleontological interpretation of species' range charts. If the method is used for such studies, the possibility of suggesting an environmental or biological cause for observed patterns, when in fact no such cause exists, will be diminished. The method is applied to a large, well-studied fossil data set to illustrate some possible species' range patterns which result only from sample size inequities. As examples, stepwise extinction and species richness patterns are predicted for situations where, in reality, none exist.

A review of the processes required for exceptional preservation of soft-bodied fossils demonstrates that anoxia does not significantly inhibit decay and emphasizes the importance of early diagenetic mineralization. Early diagenesis is the principal factor amongst the complex processes leading to soft-part preservation. The development of a particular preservational mineral is controlled by rate of burial, amount of organic detritus, and salinity. A new causative classification of soft-bodied fossil biotas is presented based upon fossil mineralogy and mineral paragenesis.

Actualistic experiments have quantified rate of anaerobic decay and associated mineralization around proteinaceous macro-organisms. Carcasses of the polychaete worm Nereis and the eumalacostracans Nephrops and Palaemon were buried in airtight glass jars filled with sediment and water from marine, brackish, and lacustrine environments. Over a period of 25 weeks the contents were examined to determine the state of decay and were chemically analyzed to monitor early diagenetic mineralization (two methods for such analysis are reviewed). Decay processes were active in the experimental conditions despite anoxia and had virtually destroyed the carcasses within 25 weeks. However, decay-rate in the sulfate-reducing marine system was greater than in the methanogenic freshwater environments. Petrological and geochemical analyses of the organic remains identified discrete layers of authigenic iron monosulfide (a pyrite precursor) on the surface of the decaying Nephrops cuticle within weeks of initiating the experiment. Chemical analysis of decomposing flesh showed a marked increase in pore-water calcium content with time.

The results clearly show that anoxia is ineffective as a long-term conservation medium in the preservation of soft-bodied fossils. However, decay-induced mineralization can be very rapid so that even a slight reduction in decay rate can lead to improved levels of fossil preservation. Traditionally, stagnation and rapid burial are considered to be the main prerequisites for the preservation of soft-bodied fossils and the formation of Konservat-Lagerstätten. Clearly these factors are only important in that they promote early diagenetic mineralization. This is the only way to halt information loss through decay.

The Late Devonian extinction event was not geologically “instantaneous,” in that extinctions during the epoch are not concentrated into a single sharp pulse at the end of the Frasnian. Extinction rates are elevated for a period of at least 2 to 4 m.y. during the middle and late phases of the Frasnian, with maximum rates occurring generally 2 m.y. before the terminal Frasnian. Neither was the Late Devonian biotic crisis a “gradual” event. In the analysis of the evolution of ecosystems, it is misleading to consider the pattern of extinction rates alone. Frasnian marine ecosystems flourished during the same time interval characterized by elevated extinction rates because origination rates of new species are higher, per time interval, than corresponding extinction rates. This pattern of relative origination/extinction rates abruptly reversed during the latest Frasnian—precipitating a rapid loss of species diversity. Within limits of current stratigraphic correlation, the ecosystem collapse appears to have occurred simultaneously in such widespread geographic regions as New York State (U.S.A.) and the southern Urals (U.S.S.R.).

In viewing the Late Devonian event from an ecological perspective, the most important question is not “What triggered the elevated extinction rates?”, but rather “What was the inhibiting factor that caused the cessation of new species originations?”

The Cretaceous–Tertiary (K–T) extinction reduced the gamma diversity of molluscs on the U.S. Gulf Coast from over 500 species in the late Maastrichtian to a little over 100 species in the early Danian. Gamma (total) diversity increased in a series of steps that generally tracked temperature, to a high of around 400 species in the late Middle Eocene, at which time diversity declined in the Late Eocene–Oligocene extinctions. The molluscan radiation occurred in at least two distinct phases: 1) an Initial Radiation Phase in which certain families underwent unusually high speciation, apparently filling ecological niches vacated by the extinction, followed by extinction of many of the species in these families in the late Danian; and, 2) a Secondary Radiation Phase where gamma diversity gradually increased and new genera gradually appeared. The fact that the gamma diversity of molluscs did not reach pre-extinction levels before the next extinction in the Late Eocene suggests that molluscan faunas may spend much of their evolutionary time recovering from these extinctions.

Trophic diversity within guilds of terrestrial predators is explored in three modern and two ancient communities. The modern communities span a range of environments including savannah, rainforest, and temperate forest. The paleocommunities are North American, Orellan (31–29 Ma), and late Hemphillian (7–6 Ma), respectively. The predator guilds are compared in terms of: 1) species richness; 2) the array of feeding types; and 3) the extent of morphological divergence among sympatric species. Feeding type is determined from dental measurements that reflect the proportion of meat, bone, and non-vertebrate foods in the diet. Measurements include estimates of canine shape, tooth size, cutting blade length, and grinding molar area. Morphological divergence among sympatric predators is measured by calculating Euclidean distances among species in a six-dimensional morphospace. Results indicate that the number of predator and prey species are roughly correlated in both ancient and modern communities. Two of the predator guilds, the late Hemphillian and modern Yellowstone, contain relatively few species and appear to be the result of extinction without replacement. Despite differences in history, age, and environment, the extent of morphological divergence within guilds does not differ significantly for the sampled communities. It is clear that the basic pattern of adaptive diversity in dental morphology among coexisting carnivores was established at least 32 million years ago. It appears that interspecific competition for food has acted similarly to produce adaptive divergence among sympatric predators in communities that differ widely in time, space, and taxonomic composition.

Substantial geographic coverage in paleontological study is essential in testing evolutionary models of phyletic gradualism and punctuated equilibrium. We present a multivariate morphometric study of the late Neogene planktonic foraminiferal clade Globoconella using specimens from four Deep Sea Drilling Project sites (DSDP 284, 207A, 208, and 588) along a latitudinal traverse in the southwest Pacific.

During the Late Miocene (7 Ma to 5 Ma), populations of the ancestral species Globorotalia (Globoconella) conomiozea formed a geographic cline showing continuous morphological variation from the temperate sites (DSDP 284 and 207A) to the warm subtropical sites (DSDP 208 and 588). Populations living to the south had higher conical angle and fewer chambers in the final whorl compared to the northern populations. Nevertheless, populations across the entire cline exhibited a coherent, directional trend towards having larger conical angle and fewer chambers through time. At the Miocene/Pliocene boundary, the intensification of the Tasman Front (Subtropical Divergence) possibly isolated the peripheral populations in the warm subtropics from the central stocks of the temperate water masses. The evolutionary trends became decoupled: the central populations gradually lost their keel and transformed into G. (G.) sphericomiozea, while the peripheral populations in the warm subtropical areas retained their keel and evolved into a flattened species, G. (G.) pliozea.

The gradual transformation of G. (G.) conomiozea terminalis (a form retaining a keel) into G. (G.) sphericomiozea (a form lacking a keel) occurred during an interval of about 0.2 m.y., with all measured morphologic variables showing continuous and steady changes. The evolution of the central populations follows the model of phyletic gradualism. In peripheral populations, the origin of the descendant species G. (G.) pliozea from the ancestor G. (G.) conomiozea terminalis occurred very rapidly within an interval of less than 0.01 m.y. The population size of G. (G.) pliozea was small at the incipient stage at about 5.05 Ma, but increased rapidly to become dominant during the next 0.2 m.y. when the ancestral species G. (G.) conomiozea terminalis became locally extinct. Following speciation, G. (G.) pliozea exhibited morphological stasis for about 0.6 m.y., until the central stock form G. (G.) puncticulata migrated back to the warm subtropics; during the next 0.5 m.y. of their sympatry, there is no sign of hybridization between these two sister species. The evolution of G. (G.) pliozea follows the model of punctuated equilibrium.

The evolution of the Globoconella clade shows both phyletic gradualism and punctuated equilibrium. These two “alternative” evolutionary models complement each other rather than being mutually exclusive. Both models are indispensable towards providing a complete picture of the evolution of Globoconella.

Cohort analysis is used to investigate survivorship of trilobites originating during the Cambrian and Ordovician. Using a time-homogeneous branching model, it is estimated that trilobite genera and species originating during the Ordovician survived three times longer than Cambrian genera and species. Monte Carlo simulation of survivorship is used to show that (1) Cambrian and Ordovician survivorship are significantly different, (2) Ordovician cohorts conform more closely to the time-homogeneous model than do Cambrian cohorts, and (3) deviations from temporal homogeneity are more often produced by extraordinary extinction than by unusually slow turnover.

When Early Ordovician cohorts are decomposed into genera within families that originated in the Cambrian versus the Ordovician, no evidence that Cambrian and Ordovician survivorship differences are clade-specific can be found. Ordovician genera of Cambrian affinity and of Ordovician affinity become extinct at similar rates.

Some of the ultimate causes of these differences in survivorship include (1) taxonomic inconsistency, (2) greater environmental stability in the Ordovician, and (3) more highly structured ecosystems in the Ordovician that may have led to the weeding out of extinction-prone taxa.

Lunules have evolved independently in several groups of clypeasteroids, including the Rotulidae, Astriclypeidae, Mellitidae and Scutasteridae. In this paper, only the monophyletic assemblage Monophorasteridae plus Mellitidae is considered. Lunules result from modifications of the growth patterns of test plates which bring about changes in relative growth in specific directions. It is unnecessary to postulate resorption of the test. Ambulacral lunules, which are known to have hydrodynamic functions, originated as part of a series of changes: 1) bifurcation of food grooves, 2) formation of pressure drainage channels, 3) lobulation of the ambitus (as in Monophoraster), 4) complete lunule formation. The anal lunule shares the hydrodynamic function but arose separately, as a developmental aberration. The position of the periproct is highly variable but it is most commonly located at the junction of suture lines, at plate corners. In Scutella it is located between the first and second post-basicoronal plates. In Monophoraster the periproct is located further forward, between the first post-basicoronal plates, and the small anal lunule occupies the junction point between first and second post-basicoronals. It is hypothesized that the anal lunule originated as an all-or-nothing event following the forward migration of the periproct and failure to resume normal plate growth at the sutural junction point between first and second post-basicoronal plates. Its walls were derived from paired interambulacral supports. The hypothesis is discussed in connection with the ontogenetic formation of the anal lunule in living mellitid sand dollars.

Because the paleontologic concept of Coccolithophoridae species is restricted and far removed from the biologic concept, which itself is not yet satisfactorily established, calcareous nannofossil taxonomy remains in an unsatisfactory state. This situation is clearly reflected by various authors' widely different interpretations of the phylogenetic relationships among species in a given genus and among genera. Examples taken from the extant genus Helicosphaera suggest that because of parallel evolution, delineation of phylogenetic relationships among coccolith morphospecies using morphologic data alone are hazardous, as is delineation of phylogenetic relationships among closely related genera.

Nipponites, a Late Cretaceous nostoceratid ammonite, shows a peculiar meandering shell growth in the middle-late stage. Assuming neutral buoyancy, and a constant aperture angle relative to the sea bottom, meandering growth of this ammonite was modeled by computer simulation. In this model, the meandering shell growth is controlled by regulation of life orientation. The remarkable similarity in the coiling modes and rib obliquity patterns between the computer-simulated and actual specimens strongly suggests a free living mode of life in Nipponites with an approximately neutral buoyancy. The simulation also suggests that morphological saltation from a simple helicoid form like Eubostrychoceras japonicum to a meandering shell form like Nipponites occurred abruptly without any intermediate form by minor change of the upper and lower limits of growth direction.

The global diversification of the class Bivalvia has historically received two conflicting interpretations. One is that a major upturn in diversification was associated with, and a consequence of, the Late Permian mass extinction. The other is that mass extinctions have had little influence and that bivalves have experienced slow but nearly steady exponential diversification through most of their history, unaffected by interactions with other clades. We find that the most likely explanation lies between these two interpretations. Through most of the Phanerozoic, the diversity of bivalves did indeed exhibit slow growth, which was not substantially altered by mass extinctions. However, the presence of “hyperexponential bursts” in diversification during the initial Ordovician radiation and following the Late Permian and Late Cretaceous mass extinctions suggests a more complex history in which a higher characteristic diversification rate was dampened through most of the Phanerozoic. The observed pattern can be accounted for with a two-phase coupled (i.e., interactive) logistic model, where one phase is treated as the “bivalves” and the other phase is treated as a hypothetical group of clades with which the “bivalves” might have interacted. Results of this analysis suggest that interactions with other taxa have substantially affected bivalve global diversity through the Phanerozoic.

Evolutionary morphologists and paleontologists have long questioned whether there are general historical patterns to the distribution of morphological types. Few studies have rigorously addressed that question. This study tests the decoupling hypothesis, which predicts an increase in the number of morphological constraints with a reduction in the number of independent elements. Eight cases of historical transformation of the epicoracoid cartilages of frogs were selected for analysis. Similar morphological shape changes occurred across the independently derived historical transformations as determined by a triangle analysis of shape. These results support the decoupling hypothesis and indicate that there may be generalized historical pathways of structural change. This finding is important for the development of a predictive theory in evolutionary morphology.