When asked how he might sum up his long life, an aged Benjamin Franklin replied that he had, at least, been useful to his fellows. Paleobiology is still young, as our journal has just completed 20 years and now begins this season of its majority, while the vibrancy of our subject certainly proclaims a vigorous incipiency. But academic generations are short, a mere 5 years or so, and Paleobiology has therefore enjoyed adequate time to demand judgment by accomplished results, not simple promise. When, fifteen years ago (6, 1, 80), I wrote a similar piece to celebrate the completion of this journal's first five volumes, I spoke in my title about “the promise of paleobiology”; now I must refer to “the task of paleobiology” and delineate what has been accomplished, and how much we have yet to do.

Two intervals of the Phanerozoic stand out as times of biosphere-scale revolution in the sense that biogeochemical cycles came under increased control by organisms. These are the early Paleozoic (extending from just before the Cambrian to the Middle Ordovician, a duration of about 100 m.y.), characterized by the appearance of predators, burrowers, and mineralized skeletons, and by the subsequent diversification of planktonic animals and suspension-feeders; and the later Mesozoic (latest Triassic to mid-Cretaceous, a duration of somewhat more than 100 m.y.), marked by a great diversification of predators and burrowers and by the rise of mineralized planktonic protists. This paper explores the economic conditions that make such revolutions possible.

I argue that opportunities for innovation and diversification are enhanced when raw materials and energy are supplied at increasing rates, or when organisms gain greater access to these commodities through rising temperatures and higher metabolic rates. Greater per capita availability of resources enables populations to grow; lessens or alters ecological constraints on functional improvement; makes possible the evolution of high metabolic rates (large incomes), which in turn permit improvement in each of several otherwise incompatible functions; and favors the establishment and spread of daughter species arising through founder speciation. Reductions in productivity reinforce adaptational constraints and may bring about extinctions.

Massive submarine volcanism, together with its associated phenomena of warming, sea-level rise, and widening of warm-weather zones, is proposed to be the chief extrinsic trigger for the Phanerozoic revolutions. The later Mesozoic was characterized by continental rifting, which accompanied massive submarine volcanic eruptions that produced large quantities of nutrients and carbon dioxide. This activity began in the Late Triassic and peaked in the mid- to Late Cretaceous. The Early Cambrian was also a time of rifting and may likewise have been marked by large-scale submarine volcanism. Continental and explosive volcanism, weathering, and upwelling are other potential means for increasing evolutionary opportunity, but their effects are either local or linked directly or indirectly with cooling. Intense chemical weathering in the Early Cambrian, however, may have contributed to the early Paleozoic revolution.

The extrinsic stimulus was greatly amplified through positive feedback by the evolution of higher metabolic rates and other means for acquiring, trading, retaining, and recycling resources more rapidly and from a wider range of environments. Because these novelties usually require a high and predictable supply of resources, their evolution is more likely when extrinsically controlled supplies increase rather than when per capita availability is low.

In the view adopted here, the microevolutionary and microeconomic market forces of competition and natural selection operate against a backdrop of macroeconomic supply and demand. Resources are under both extrinsic and intrinsic control. Positive and negative feedbacks link processes at the micro- and macroeconomic levels. This view complements the genealogical and hierarchical conception of evolution by emphasizing that the pattern of descent is influenced by resources and by market forces operating at all scales of space and time.

The evolution of higher taxa among early Paleozoic gastropods is similar to that among early metazoans as a whole, as higher taxa diversified rapidly and early. There are two issues pertinent to this pattern. First, were greater morphologic changes concentrated in the early phases of evolution? Second, does the pattern better fit models of increasing phylogenetic constraints or increasing ecologic restrictions? This paper presents a phylogeny-based method designed to test whether amounts of morphologic evolution decreased over time. It also explores whether the data better fits models of increasing phylogenetic (i.e., developmental or genetic) constraint or increasing ecologic restriction. Two metrics of morphologic separation (i.e., the morphologic difference between sister-species) are used: (1) Euclidean distance in morphospace and (2) transition magnitude. The latter metric is calculated by a multivariate analysis of sister-species contrasts, which determines both types and magnitudes of morphologic transitions. The advantage of using transition magnitudes is that it balances the effects of transitions that either affect more morphometric characters or occur more frequently. Both metrics indicate that larger morphologic separations between sister-species were concentrated early in gastropod evolution. Among gastropods, gross shell morphology often reflects basic trophic strategy and function whereas basic internal anatomy does not. Transition magnitudes can be broken down into transitions associated with differences in basic trophic strategies and shell functional biology (“external”), and those associated with differences in basic internal anatomy (“internal”). Internal transition magnitudes show a highly significant decrease over time (p < 10–04) whereas external transition magnitudes show a much less significant decrease over time (p < 0.10) and no significant decrease after the earliest Ordovician (p ≅ 0.50). The results therefore suggest that increasing phylogenetic constraints played a greater role in the early evolution of gastropods than did increasing ecologic ones.

More than 5000 measurements were taken on over 1000 specimens of two species of brachiopods, Mediospirifer audaculus and Athyris spiriferoides, from the Middle Devonian Hamilton Group of New York state. Statistical analyses were performed on these data, with specimens partitioned by their occurrence in one of many paleoenvironments and stratigraphic horizons. Neither species showed substantial morphological departures between first appearance and extinction (the range of the Hamilton Group, roughly 5 m.y.). However, oscillations in morphology were discovered in both taxa.

For the two species we studied, groups of organisms occurring in a single paleoenvironment undergo moderate morphological change through time; however, the net sum of changes through time in all paleoenvironments in which these species occur is essentially zero. Therefore, stasis may be partly a property of the organization of species into different environmental populations. Different “environmental populations” may evolve, but they will typically do so in several different “directions,” generally producing no net change. The difference between the morphology of species in different environments over the whole interval of the Hamilton Group is also nil, thereby ruling out any major role that ecophenotypic effects could play in the patterns recognized herein.

Cladograms predict the order in which fossil taxa appeared and, thus, make predictions about general patterns in the stratigraphic record. Inconsistencies between cladistic predictions and the observed stratigraphic record reflect either inadequate sampling of a clade's species, incomplete estimates of stratigraphic ranges, or homoplasy producing an incorrect phylogenetic hypothesis. A method presented in this paper attempts to separate the effects of homoplasy from the effects of inadequate sampling. Sampling densities of individual species are used to calculate confidence intervals on their stratigraphic ranges. The method uses these confidence intervals to test the order of branching predicted by a cladogram. The Lophospiridae (“Archaeogastropoda”) of the Ordovician provide a useful test group because the clade has a good fossil record and it produced species over a long time. Confidence intervals reject several cladistic hypotheses that postulate improbable “ghost lineages.” Other hypotheses are acceptable only with explicit ancestor-descendant relationships. The accepted cladogram is the shortest one that stratigraphic data cannot reject. The results caution against evaluating phylogenetic hypotheses of fossil taxa without considering both stratigraphic data and the possible presence of ancestral species, as both factors can affect interpretations of a clade's evolutionary dynamics and its patterns of morphologic evolution.

Scleractinian corals first appeared during Triassic time in tropical shallow water environments. Controversy surrounds the paleoecology of scleractinian corals of the Late Triassic. Were they like their living counterparts, capable of supporting reefs, or had they not yet coevolved the important association with zooxanthellae that facilitated reef growth and construction? Indirect evidence suggests that some Upper Triassic corals from the Tethys played important constructional roles as reef builders within tropical carbonate complexes of the Tethys. To evaluate this idea, we have employed a geochemical approach based on isotope fractionation to ascertain if Late Triassic corals once possessed zooxanthellae.

We have determined evidence for the ancient presence of algal symbiosis in 13 species of Triassic scleratinians from reef complexes in Turkey and northern Italy. In contrast, two higher latitude Jurassic species used as a control group for isotope analysis, lacked isotopic indications of symbiosis. These findings, together with stratigraphic and paleoecologic criteria, support the contention that Late Triassic scleractinian corals inhabiting shallow-water carbonate complexes of the Tethys were predominantly zooxanthellate, like their living counterparts from present day reefs.

We view the zooxanthellate condition in calcifying reef organisms as a necessary prerequisite for constructional reef development. Our results emphasize the power of stable isotope studies in helping to answer paleobiological questions.

Several metrics, including average difference among species, range of occupied morphological space, and number of character-state combinations, are used to investigate morphological diversification in Paleozoic crinoids. Despite several phases of taxonomic diversification, the maximal level of disparity reached in the Ordovician remained essentially unsurpassed. Although new regions in morphological space were occupied after the Devonian, these were not as extensive as those that had been evacuated prior to the Carboniferous. This discordance between extensive total morphological change and limited net change further supports previous arguments for the importance of morphological constraints in crinoid evolution. Major changes in the occupation of morphological space correspond with changes in taxonomic diversity within certain higher taxa. The extent to which advanced cladids (Poteriocrinina) appear to expand into new morphological space is exaggerated by the large number of very similar species in this group. If fewer species are sampled, by considering only those forms that differ from each other by at least some prescribed amount, poteriocrines appear to be less extreme morphologically. In contrast, other groups that seem to occupy unique regions in morphological space continue to do so even if fewer of them are sampled. Major crinoid clades—Camerata and Cladida+Flexibilia—do not show the same evolutionary pattern as Crinoidea, but instead exhibit a more gradual diversification of morphology. This observation provides additional support for the existence of qualitative differences among taxa of different rank.

Paleobiologists have used taxonomic data for several types of diversity studies. Some systematists have charged that this practice obfuscates actual historical patterns of clades because many traditionally defined higher taxa are not monophyletic. Some have questioned whether ranked taxa ever represent comparable units, even when monophyletic. This study contrasts diversity patterns implied by phylogenetic estimates with those implied by ranked taxa. Early Paleozoic gastropods are useful as a test case because their generic taxonomy does not reflect the phylogenetic systematic philosophy, and fewer than one third of the genera represent monophyletic clades. Phylogenetic diversity is described in two ways: (1) numbers of lineages (i.e., observed plus phylogenetically implied “ghost lineages”), and (2) numbers of monophyla (i.e., clades whose sister taxa are other clades rather than species). “Monophyla” as tallied here are monophyletic relative to their contemporaries and older clades; however, they can be paraphyletic relative to “future” monophyla. Phylogenetic diversity is tallied with both maximum and minimum “ghost lineage” interpolations in order to reflect different possible speciation patterns and timings of speciation. Phylogenetic diversity as implied by a stricter cladistic criterion (i.e., taxa that are monophyletic relative to their contemporaries, older taxa and younger taxa) is discussed also.

First differences between substage-to-substage standing diversities reveal significant congruence between generic data and both types of phylogenetic data. Taxonomic and phylogenetic data imply a major extinction event at the end of the Ordovician, although the phylogenetic data suggest greater extinction levels than do the taxonomic data. Both data sets also suggest diversity-dependent diversification reminiscent of logistic growth, which is the pattern predicted if one or a few major ecologic factors were constraining the diversification of gastropods. However, diversity described by strict Hennigian taxa is not highly congruent with diversity as described by either lineages or monophyla. Comparing subclade dynamics requires extensive redefinition of traditional orders, but lineages, monophyla and genera all suggest that the two major subclades had different logistic diversification patterns, with one (“murchisonioids”) having a higher K than the other (“euomphaloids”). The concern that phylogenetic and taxonomic data might imply very different evolutionary histories is not borne out by gastropods, despite the nonphylogenetic nature of their traditional taxonomy.

Speciation processes are only rarely studied with fossil materials, even though in principle hypotheses of speciation patterns are most directly testable in the fossil record. We quantitatively document in two widely separated South Pacific DSDP holes the mid-Pliocene speciation of the planktonic foraminifer Globorotalia truncatulinoides. Speciation, with continuous geographic co-occurrence of ancestor and descendant forms, occurred simultaneously at both localities over a period of ~500,000 years. This suggests a sympatric speciation process that involved a large, geographically extensive population. Globorotalia truncatulinoides underwent its most rapid and extensive evolutionary change between ~2.8 and 2.5 Ma. This time interval corresponds to the development of northern hemisphere glaciation, suggesting that climate-controlled paleoceanographic change may have played a significant role in the evolution of G. truncatulinoides.

Approximately half of the Caribbean Oligocene reef coral fauna became locally extinct during the Early Miocene; roughly two thirds of the genera driven to local extinction still survive in the Indo-Pacific. Coral genera with lecithotrophic larvae (brooders) preferentially survived, over those with planktotrophic larvae (broadcasters). Among 37 genera for which we inferred reproductive mode, 73% of brooding genera survived the Oligocene/Miocene extinction events, while only 29% of the broadcasting genera survived. The proportion of brooders to broadcasters also increased markedly. During the late Oligocene, 47% of Caribbean reef coral genera were broadcasters, but in the middle Miocene, only 32% of the genera were broadcasters.

Survivorship in Puerto Rican reefs was correlated with tolerance of cold and turbid conditions. Genera tolerant of both cold water and turbidity had much higher survival rates than those tolerant of turbidity alone. Only 25% of the genera that could tolerate neither cold water nor turbidity survived. Most of the eurytopic genera were brooders, while most of the stenotypic genera were broadcasters.

We present two hypotheses that may account for the preferential survivorship of brooders: the recruitment hypothesis, and the dispersal hypothesis. The recruitment hypothesis holds that brooders survive preferentially because lecithotrophic larvae have higher recruitment success than do planktotrophic larvae in marginal habitats, such as upwelling zones. This is supported by the correlation of brooding and eurytopy. The dispersal hypothesis suggests that brooders survive preferentially because lecithotrophic larvae, which typically inherit zooxanthellae from the egg, have a longer larval lifespan and, hence, a wider potential dispersal range, than planktotrophic larvae, which typically capture zooxanthellae from the water column. Biogeographic range data, however, do not support this second hypothesis: modern Indo-Pacific brooding and broadcasting genera have nearly identical ranges, and many brooding species have narrower longitudinal ranges than do broadcasting species. Preferential survivorship of brooding corals contrasts sharply with survivorship patterns among molluscs during extinction events; among molluscs, broadcasters are favored over brooders.

A major increase in upwelling at the Oligocene/Miocene boundary was probably responsible for this extinction/geographic restriction event. Preferential survival of brooding and mixed mode coral genera appears to be a product of their being better able to recruit and survive in marginal conditions such as upwelling zones.

A hierarchical branching model is presented that tracks diversity of lineages and paraclades simultaneously. It is based on three parameters: (1) lineage origination within a paraclade, (2) lineage origination that founds new paraclades, and (3) lineage extinction. The model assumes stochastic constancy of evolutionary rates and, therefore, exponential growth of diversity is the null expectation. The probabilistic model can be used to generate confidence intervals for exponential growth through Monte Carlo simulation. These confidence intervals can be compared to empirical curves to test the null hypothesis of exponential growth. The analytic solution for paraclade diversity gives the statistical expectation for the probabilistic model. It can be applied to empirical diversity curves of supraspecific taxa to estimate total speciation rate, the intrinsic rate of increase, and the proportion of speciation events that found new paraclades. Two factors were identified with the model that may cause diversity curves for higher taxa (e.g., genera, families) to deviate from simple exponential growth during initial phases of diversification: (1) stochastic fluctuation at low diversity, and (2) taxonomic structure (i.e., species/genus ratio). These sources of variability should be evaluated before macroevolutionary explanations for specific diversity trends are invoked. Two kinds of evolutionary radiations were investigated with this model: (1) prolonged periods of low diversity (macroevolutionary lag) in cheilostome bryozoans and mammals prior to their main phases of diversification, and (2) rapid bursts in diversity of bivalves following the late Permian and end-Cretaceous mass extinctions. In all cases these patterns were found to deviate significantly from the null expectation of exponential growth lending support to previous macroevolutionary explanations. Finally, rates of speciation during radiations were compared to rates of speciation during background times for articulate brachiopods, cheilostome bryozoans, bivalves, and mammals. Speciation events that found new paraclades tend to make up a larger proportion of total speciation events during radiations compared to background times indicating that the opportunity for new adaptive zones to be filled at higher taxonomic levels is proportionately higher during these periods of increased evolutionary activity, and is not simply a result of an increased frequency of speciation.

We analyze a new compilation of Neogene to Recent (22-0 Ma) Caribbean coral occurrences to determine how ecological and life history traits at the population level affect long-term evolutionary patterns. The compilation consists of occurrences of 175 species and 49 genera in one continuous (> 5 m.y.) sequence and 22 scattered sites across the Caribbean region. Previous study of evolutionary rates using these data has shown that both extinction and origination were accelerated between 4 and 1 Ma, resulting in large-scale faunal turnover. Categories for three morphological and two reproductive variables (colony size, colony shape, and corallite size; and sex, and mode of embryonic development; respectively) are assigned to each species in the compilation. Comparisons of the ecological variables with evolutionary rates using randomization procedures and modified analysis of variance show that only colony size was strongly related to rates of extinction and origination during either normal background times or times of accelerated extinction. Extinction rates were lower in species with large colonies, because species with small massive colonies tend to live in small, short-lived populations with highly fluctuating recruitment rates. During turnover, extinction rates increased disproportionately in species with small colonies. Origination rates are found to be less related to ecological variables, although species with small massive colonies originated at higher rates prior to turnover.

Accelerated turnover may have therefore involved an increase in local population extinction rates that caused increased rates of both species extinction and origination across the entire fauna. Since extinction rates accelerated disproportionately with respect to colony size, the overall result was a relative increase in species with large colonies. After severe disturbance, one might expect that populations of species with large colonies and high rates of fragmentation would be more likely to escape extinction, because of larger population sizes, longer generation times, and more constant rates of population increase. The modern Caribbean reef-coral fauna is therefore structured by large, long-lived colonies that are robust to regional environmental change. Many of the very taxa that allowed reef communities to escape collapse in the past are declining today in response to anthropogenic disturbances, suggesting that Caribbean reef communities may be less resilient in the future in response to ongoing environmental perturbations.

Documenting past environmental disturbances will provide a very incomplete explanation of extinctions until more data on intrinsic (e.g., phylogenetic) responses to disturbances are collected. Taxonomic selectivity can be used to infer phylogenetic inheritance of extinction-biasing traits. Selectivity patterns among higher taxa, such as between mammals and bivalves, are well documented. Selectivity patterns among lower taxa (genus, species) have great potential for understanding the dynamics underlying higher taxic turnover. Two echinoid data sets, of fossil and living taxa, indicate that species extinctions do not occur randomly within genera. Reverse rarefaction estimates of past species extinction rates assume random species extinction within higher taxa, so these widely cited extinction estimates may be inaccurate. Revised estimates based on a simulated curve imply that past species extinctions rates may be 6%–15% lower than previously cited. Possible causes for the observed selectivity patterns are discussed. These include nonrandom phylogenetic nesting of species with traits often cited as enhancing extinction vulnerability, into certain taxa. Such traits include low abundance, large body size, narrow niche breadth, and many others. Phylogenetic nesting of extinction-biasing traits at many taxonomic levels does not predict that a dichotomy of mass-background selectivity based on a few traits will occur. Instead, it predicts patterns of selectivity at many taxonomic levels, and at many spatio-temporal scales.

In phytogeographic data sets, the number of assemblages or floras from each interval may provide a test of the influence of sampling intensity on land-plant diversity. Using a data set of Silurian and Devonian compression-impression plant genera from Laurussia and the Acadian terrain, regression of five measures of land-plant diversity (total diversity, mean genus richness of floras, median assemblage diversity, most diverse assemblage, and standing diversity at interval boundaries) against the number assemblages or floras from thirteen intervals suggests that sampling bias influences all of the diversity measures to some extent, including within-habitat measures. The standing diversity of land plants at interval boundaries, the measure least influenced by sampling (r = 0.65, p = 0.05), increased steadily from the Middle Silurian to the late Givetian/early–middle Frasnian boundary, fell sharply in the early–middle Frasnian and remained low throughout the late Frasnian–middle Famennian. Standing diversity rose dramatically in the late Famennian and Strunian (latest Devonian): the Frasnian–Famennian extinction event may have affected land plants. The standing diversity of Silurian and Devonian microspore genera at interval boundaries mirrors that of compression-impression genera: neither record supports a land-plant diversity equilibrium during the Devonian.

Brachiopods generally have not been considered to be typical or significant faunal components of modern or ancient hydrothermal vent and cold-seep settings. The Early Cretaceous (Neocomian) rhynchonellide brachiopod Peregrinella has long been viewed as a paleontological curiosity because of its distinctive morphology, status as the largest Mesozoic brachiopod, anomalous stratigraphic associations, and widespread, yet discontinuous paleogeographic distribution. Examination of all worldwide Peregrinella occurrences (14) indicates restriction of this brachiopod to ancient cold-seeps. It is probable that Peregrinella grew to large sizes in such great abundances at fossil cold-seep sites because of a richly organic food supply generated by localized fluid seepage and bacterial chemosynthetic activity. Living brachiopods are not known to harbor chemosymbiotic bacteria in their tissues; however, direct chemoautotrophic utilization of reduced fluids by Peregrinella cannot be rejected or demonstrated at present. Peregrinella occurs at widely separated cold-seeps of Neocomian age (e.g., California, Mexico, Tibet, Europe), yet its mode of dispersal and larval development is unknown. In modern hydrothermal vents of the deep-sea, organism dispersal occurs along oceanic ridges, where benthic faunas display both planktotrophic and nonplanktotrophic larval-mode types. Peregrinella may represent a Mesozoic relic of a long-lived “lineage” of vent-seep associated rhynchonellides from the Paleozoic (e.g., ?Eoperegrinella, Dzieduszyckia), but major gaps in the stratigraphic record between these rhynchonellide occurrences, and the lack of rigorous phylogenetic analysis for these groups preclude a clear resolution of the origin(s) of vent-seep brachiopods at present.

Documentation of morphologic variation within and among fossil species (and larger clades) provides fundamental data needed for studies of evolution, paleoecology, and the systematic foundation required for most fields of paleobiology. In paleontological (and, frequently, biological) studies, morphologic variation is used as a general proxy for genetic variation. Although the occurrence of ecophenotypic variation is well appreciated in these studies, it is only with the use of colonial (clonal) organisms that the scope and significance of phenotypic variation can be evaluated directly. Systematic evaluation of intracolonial morphologic variation (transects through growth series) can yield insights about ecophenotypic variation in bryozoans and suggest the most appropriate methods for data collection in paleobiologic and taxonomic studies.

In this study, morphological conservatism is documented within local segments of bryozoan colonies; each zooid is generally more similar to adjacent zooids than to distant zooids within the same colony. One region of a colony, therefore, can be more similar to a region of a different colony than to a distant region of its own colony. Variation within one colony does not, however, represent the total variation among a group of specimens, indicating a colonial level of morphologic control (genetic or macroenvironmental) over morphogenesis. Directional morphogenetic gradients (associated with successive ontogenetic histories) are not recognized in these specimens, but fluctuating trends within colonies (some cyclic), were observed and are indicative of changing microenvironmental influence during skeletal formation. In order to best document morphologic variation within a population, for any type of paleobiological study, individual measurements should be widely distributed over large colony fragments and (or) a minimal number of measurements collected from each of a large number of smaller fragments.

Direct extrapolation of these results to non-colonial organisms is not appropriate at this time. However, additional, related studies with bryozoans and other colonial organisms (e.g., corals, graptolites), should provide a greater, general appreciation of relationships between morphology and genetics.

Despite exhaustive investigation of present-day Nautilus, the phylogenetic relationships of the five or six recognized species within this genus remain unclear. Mitochondrial and nuclear DNA sequence data plus a suite of morphological characters are used to investigate phylogenetic relationships. Systematic analysis of the morphological variation fails to characterize described species as independent lineages. However, DNA sequence analysis indicates that there are three geographically distinct clades consisting of western Pacific, eastern Australian/Papua-New Guinean, and western Australian/Indonesian forms. The morphologically and genetically distinct species Nautilus scrobiculatus falls outside the three geographically recognized assemblages. Members of the genus Nautilus also exhibit low levels of sequence divergence. All these data suggest that Nautilus is currently undergoing diversification, which may have begun only several million years ago. These data also suggest that some of the morphological features used to define Nautilus species may simply represent fixed variations in isolated populations within the same species.