Ecologists attempt to explain species diversity within Recent seabird communities in terms of Recent oceanographic and ecological phenomena. However, many of the principal oceanographic processes that are thought to structure Recent seabird systems are functions of geological processes operating at many temporal and spatial scales. For example, major oceanic currents, such as the North Pacific Gyre, are functions of the relative positions of continents and Antarctic glaciation, whereas regional air masses, submarine topography, and coastline shape affect local processes such as upwelling. I hypothesize that the long-term development of these abiotic processes has influenced the relative diversity and community composition of North Pacific seabirds. To explore this hypothesis, I divided the history of North Pacific seabirds into seven intervals of time. Using published descriptions, I summarized the tectonic and oceanographic events that occurred during each of these time intervals, and related changes in species diversity to changes in the physical environment. Over the past 95 years, at least 94 species of fossil seabirds have been described from marine deposits of the North Pacific. Most of these species are from Middle Miocene through Pliocene (16.0-1.6 Ma) sediments of southern California, although species from Eocene to Early Miocene (52.0-22.0 Ma) deposits are from Japan, British Columbia, Washington, and Oregon. During the history of the North Pacific seabirds, thee were many widespread (global) and local (California Current) oceanographic events, but the underlying physical processes that affected the diversity of seabirds throughout the North Pacific were tectonic changes that led to sequential stages of thermal isolation and refrigeration of Antarctica. Besides these broad-scale phenomena, the uplift of the Isthmus of Panama and the initiation and intensification of coastal upwelling in the California Current directly affected the diversity of seabirds in southern California. For the most part, Middle to Late Miocene and Pliocene seabird faunas from California resembled Recent communities, but with several exceptions: (1) the extreme diversity and abundance, but subsequent extinction of gannets (Morus) and flightless alcids (mancallids); (2) the comparatively low abundance and diversity of cormorants and shags (Phalacrocoracidae) until the Late Pliocene; and (3) the absence of marine Laridae until the Late Pliocene. By affecting both the physical and biological environments, geological factors such as Antarctic glaciation, eustatic changes in sea levels, and local tectonic activities influenced and will continue to influence the structure of seabird systems in the North Pacific.

The extreme morphological simplicity of lingulide brachiopod shells makes them particularly useful for investigating the species-level taxonomic resolution of the fossil record as well as the relationships between taxonomy, morphological complexity, and evolutionary rates. Lingulides have undergone little change in shell morphology and have had low taxonomic diversity since the Paleozoic. Is this pattern an evolutionary phenomenon or an artifact of the shell's simplicity? Multivariate methods were used to establish morphogroups among seven populations of four extant species of Glottidia. Six characters (three shell dimensions and three internal septa) were measured for 162 specimens from field and museum collections. All populations follow similar allometric trajectories: internal septa display positive allometry and shell dimensions display negative allometry. The allometric pattern may reflect D'Arcy Thompson's Principle of Similitude. Principal component analysis does not reveal any distinct clusters in Glottidia morphospace but suggests that some differences independent from ontogeny exist among the populations. Size-free canonical variate analysis indicates the presence of five size-invariant groups that are statistically distinct. Bootstrap-corrected error rates indicate that four specimens are enough to classify a sample correctly at α = 0.05 and eight specimens at α = 0.01. The groups are consistent with neontological classification with the exception of two populations of G. pyramidata identified by discriminant analysis as two distinct groups. The size-free morphogroups reflect geographic separation rather than ontogenetic or substrate differences among the populations.

Despite the morphological simplicity of the shell, size-free multivariate analysis of Glottidia delineates groups that offer taxonomic resolution comparable with the neontological classification. The method offers a promising tool for identifying natural morphogroups on the basis of few morphological characters. Moreover, the agreement between neontological taxonomy and the morphogroups suggests that the size-free approach can be applicable for evaluating the reality of the low diversity and turnover rates observed in the fossil record of lingulide brachiopods (= Family Lingulidae). Assuming that the neontological species of Glottidia are biologically meaningful, this study shows that morphological simplicity of lingulides does not necessarily result in taxonomic underresolution. Our analysis, as well as several previous case studies, suggests that taxonomic diversity and turnover rates do not have to be dependent on the morphological complexity of preservable parts. In many cases, when rigorous quantitative methods are employed, the differences in the rates of morphological evolution may be a real evolutionary phenomenon and not artifacts of morphological complexity.

Recent molecular phylogenetic and molecular clock data both suggest a pre-Mesozoic age for the divergence of the angiosperm lineage from other seed plants, greatly predating the confirmed fossil record of the angiosperm crown group. In addition, molecular phylogenetic studies have not supported the morphologically based conclusion that gnetophytes are the extant sister group to angiosperms. We examine these relationships and divergence ages by using a novel approach of examining the presence of oleanane. This includes the development of methods using zeolites to preferentially reduce hopanes that can co-elute with oleanane. The presence of this molecular fossil strongly correlates with angiosperm diversification; in its functionalized form, along with its triterpenoid precursors, it is found in many living angiosperms. Our data show that among non-angiosperm seed plants examined thus far, oleanane is found only in fossil Cretaceous Bennettitales and Permian Gigantopteridales, both of which share characteristics with angiosperms. Previous morphological phylogenetic results indicate Bennettitales could be a sister group to or member of the angiosperm stem lineage, and results of our preliminary phylogenetic analysis including the Gigantopteridales suggests the same. Our data, based on a new pyrolysis method to treat living species, support previous research indicating that oleanane and its precursors are absent in living gnetophytes. If oleanane originated once in seed plants then the angiosperm stem lineage would have diverged from other seed plant lineages by the late Paleozoic.

The megafauna and associated behavioral traces of two deep-sea benthic environments, the central Arctic and Antarctic, with a surface primary productivity differential of 104 were compared to assess the role of food availability in foraging strategy and community structure. Bottom photographs, analyzed for megafauna and trace density and diversity at comparable depths in the Arctic Canada Basin and the Antarctic Bellingshausen Basin, indicated that trace frequency was inversely proportional to organism density but that trace diversity directly reflected organism diversity. Those traces identified in the fossil record to represent efficient foraging strategies, i.e., the Nereites facies, were conspicuously absent at all depths in the Arctic and present at all depths in the Antarctic, in contradiction of the paradigm of increasing behavioral complexity and sediment exploitation as food availability decreases. Presence or absence of surface-grazing organisms seems to exert a greater influence on trace diversity than depth or nutrient supply. Trace density, however, may reflect episodic sedimentation events which intermittently influence the deep-sea trophic regime.

It is commonly believed that the brains of the ancestors of modern carnivores (miacids) were superior to (e.g., larger than) those of other early carnivores (creodonts and mesonychids). Examination of the fossil record of brains of early carnivores reveals no evidence to support that belief. Moreover, evolutionary trends towards increasing relative brain size and an expansion of neocortex are seen in both miacids and creodonts. The neocortex expanded in a different way in miacids than in creodonts and mesonychids (evidenced by different sulcal patterns), but the biological significance of the observed differences is unknown.

The end-Permian mass extinction is commonly portrayed not only as a massive biodiversity crisis but also as the time when marine benthic faunas changed from the Paleozoic Fauna, dominated by rhynchonelliform brachiopod taxa, to the Modern Fauna, dominated by gastropod and bivalve taxa. After the end-Permian mass extinction, scenarios involving the Mesozoic Marine Revolution portray a steady increase in numerical dominance by these benthic molluscs as largely due to the evolutionary effects of an “arms race.” We report here a new global paleoecological database from study of shell beds that shows a dramatic geologically sudden earliest Triassic takeover by bivalves as numerical dominants in level-bottom benthic marine communities, which continued through the Early Triassic. Three bivalve genera were responsible for this switch, none of which has any particular morphological features to distinguish it from many typical Paleozoic bivalve genera. The numerical success of these Early Triassic bivalves cannot be attributed to any of the well-known morphological evolutionary innovations of post-Paleozoic bivalves that characterize the Mesozoic Marine Revolution. Rather, their ability to mount this takeover most likely was due to the large extinction of rhynchonelliform brachiopods during the end-Permian mass extinction and aided by their environmental distribution and physiological characteristics that enabled them to thrive during periods of oceanic and atmospheric stress during the Permian/Triassic transition.

Discovery of several new specimens of the gigantic Eocene ground bird Diatryma gigantea from the Willwood Formation of northwestern Wyoming, has prompted an analysis of its feeding apparatus and an assessment of the mode of life of this unusual bird. Diatryma exhibits many of the features predicted by biomechanical models to occur in animals delivering large dorsoventral bite forces. Similarly, the mandible of Diatryma, which was modeled as a curved beam, appears well equipped to withstand such forces, especially if they were applied asymmetrically. Interpretation of these size-independent biomechanical properties in light of the large absolute skull size of Diatryma suggests a formidable feeding apparatus. The absence of modern analogues complicates the determination of just how this unique skull morphology correlates with diet. Suggestions that Diatryma was an herbivore seem improbable in that they require the postulation of excessively high safety factors in the construction of the skull. The traditional hypothesis of Diatryma as a carnivorous bird accords as well or better with the data at hand. Carnivory raises the probability of “accidental” encounter with bones, thus explaining the high safety factors. In fact, the skull and mandible of Diatryma are so massive that bone crushing may have been an important behavior. Diatryma could have been a scavenger. However, limb allometry and phylogenetic interpretation of limb proportions call into question the picture of Diatryma as a slow, plodding graviportal animal, suggesting that active predation was within its behavioral repertoire.

The hyrax, a living primitive subungulate, makes an ideal model in terms of its molar morphology for early lophodont ungulates. Cine fluoroscopic studies of mastication show that the proposed Phase II in jaw movement, experimentally demonstrated in primates, but surmised in ungulates only from molar wear facets, does in fact exist in this animal, but does not appear to be an important component of the masticatory cycle. Comparison of tooth wear in early Tertiary ungulates with contemporaneous primates shows that, whereas primates enlarge the Phase II wear facets on the molars as they move into more herbivorous niches, the same is not true for ungulates, and in fact Phase II may be lost altogether in grazing forms. It is concluded that “grinding” in herbivorous ungulates is a process fundamentally different from that described for herbivorous primates.

Cantius is an early Eocene adapid primate with an exceptionally well known fossil record. Measurements were recorded to describe the size and shape of upper and lower first molars collected from a measured stratigraphic section in the Clarks Fork Basin of Wyoming. Rates of change of size and shape are quantified by calculating evolutionary rates in standard deviation units per generation (haldanes). Temporal scaling of rates shows that change in size was generated by a significantly nonrandom directional process, while change in shape was generated by a significantly nonrandom stabilizing process. Size change in Cantius is interpreted to be the result of weak directional selection, and shape change is interpreted to be the result of strong stabilizing selection.

Diversity dynamics of the Permian–Triassic land plants in South China are studied by analyzing paleobotanical data. Our results indicate that the total diversity of land-plant megafossil genera and species across the Permian/Triassic boundary (PTB) of South China underwent a progressive decline from the early Late Permian (Wuchiapingian) to the Early-Middle Triassic. In contrast, the diversity of land-plant microfossil genera exhibited only a small fluctuation across the PTB of South China, showing an increase at the PTB. Overall, land plants across the PTB of South China show a greater stability in diversity dynamics than marine faunas. The highest extinction rate (90.91%) and the lowest origination rate (18.18%) of land-plant megafossil genera occurred at the early Early Triassic (Induan), but the temporal duration of the higher genus extinction rates (>60%) in land plants was about 23.4 Myr, from the Wuchiapingian to the early Middle Triassic (Anisian), which is longer than that of the coeval marine faunas (3–11 Myr). Moreover, the change of genus turnover rates in land-plant megafossils steadily fluctuated from the late Early Permian to the Late Triassic. More stable diversity and turnover rate as well as longer extinction duration suggest that land plants near the PTB of South China may have been involved in a gradual floral reorganization and evolutionary replacement rather than a mass extinction like those in the coeval marine faunas.

The most important topics discussed thus far by community paleoecologists are: community approach to paleoenvironmental reconstruction, community development in evolutionary time, and community constraints upon species evolution. The first step in community-paleoecological analysis is to reconstruct the paleocommunity or paleoecosystem. However there are severe methodological limitations to any inference drawn from the composition and structure of a fossil assemblage. These result from various taphonomic biases. These constraints upon reliability of a community-paleoecological analysis are the least severe in the case of Cenozoic (and possibly Cretaceous) tropical to subtropical, molluskdominated, subtidal, benthic communities.

The basic assumption of community paleoecology is that communities or biocoenoses represent a distinct, real level of biotic organization achieved through ecological integration of and coevolution among the species. This assumption seems to be invalid for two reasons. (1) The actual degree of community integration is in general insufficient to induce any driving forces for a structural development as predicted by the system theory. (2) The concept of biological reality and distinctness of the community level of biotic organization implies assignment of a significant role to group selection. The assumption that ecological communities achieve with time an equilibrium state representing an optimum habitat partitioning among the component species is invalid, at least as a generalization. This idea is largely falsified and refuted by a variety of ecological studies. Ecological communities are then merely an epiphenomenon of the overlap in distributional patterns of various organisms. There is no intrinsic, biotic mechanism inducing community dynamics in either ecological, or evolutionary time.

In spite of this conclusion, the so-called “community approach” under favorable taphonomic conditions is among the most reliable methods of paleoenvironmental reconstruction, and the data on so-called “community evolution” are relevant to the problem of a relationship between actually realized niche patterns and their environmental framework. The latter problem can also be approached through analysis of longevity-frequency-distributions of chronospecies found living together in various habitats. A preliminary investigation may indicate that the precondition to optimization of niche dimensions through coevolution among ecologically related species was met in subtidal benthic habitats but not in pelagic ones. To account for a niche subdivision among planktonic organisms, one has therefore to invoke a stochastic pattern of speciation rather than coevolutionary mechanisms.

We have known since Trueman's classic work of 1922 that the Lower Jurassic Gryphaea of Britain exhibit phyletic size increase and heterochronic change in shape. Since Hallam's revisionary work in the 1960s, we have recognized that pronounced and generalized juvenilization of form accompanied this increasing size. This extensive literature provides invertebrate paleontology's most famous example of a biometrically documented, continuous anagenetic trend within a discrete lineage. But Gryphaea has also provoked great frustration because a key datum, required for a full solution, had been theoretically recognized but practically unavailable. We could identify the evolution of shape as paedomorphic, but could not specify the mode of heterochrony for this paedomorphic result because we could not standardize samples by common age or developmental stage.

In this paper, we provide sclerochronological data on sizes and shapes at specified ages marked by annual growth bands in two Jurassic sequences of Gryphaea: the classic Lower Jurassic series showing phyletic size increase with paedomorphosis, and an independent Middle–Upper Jurassic series illustrating neither size increase nor heterochrony. We prove that size increase in the classic series occurs entirely by faster growth (larger descendant sizes at the same ages as ancestors), and not by extended age (for descendants lived no longer than ancestors). The well-marked paedomorphosis of form probably arose as a correlated consequence of growing larger by extending and maintaining rapid juvenile growth rates—thus marking the heterochronic mode as a case of neoteny. The independent upper sequence, acting as a different replicate in a natural experiment, shows neither size increase nor heterochrony but does exhibit (in contrast with the classic sequence) evolution toward greater longevity.

Hallam's flow tank experiments indicated a strong adaptive advantage in shell stability for both larger size and paedomorphic form. Neotenous development provides an evolutionary pathway to the simultaneous acquisition of both favored traits—thus showing that “constraints” due to “correlations of growth” (Darwin's own phrase for the phenomenon) may be positive in promoting joint evolutionary advantages, and not only neutral (in carrying spandrels along with primary adaptations), or negative (by imposing inadaptive “baggage” upon trends in form through developmental correlation with selected traits).

Field and laboratory data indicate that all four species of Thais (Gastropoda, Muricacea) from rocky shores of the northeastern Pacific selectively attack barnacles at the margins of parietal (lateral) and opercular plates. Attacks are also more likely to be successful at plate margins. Such preferential attack and differential attack success may account for the evolutionary reduction in the number of parietal plates exhibited within seven of the eight families of balanomorph barnacles. Additional evidence suggests that predation by drilling gastropods may have also favored the evolution of strong external ribbing on these plates within some balanomorph lineages.

Radiation and plate reduction in the Balanomorpha occurred in concert with extensive radiation in muricacean gastropods, starting in the Late Cretaceous. Of the three skeletal conformations exhibited by open-surface dwelling barnacles (8, 6 and 4 parietal plates), only genera of the four-plated form have become proportionally more common in the Recent. Further, in a rather striking evolutionary experiment, a lineage of barnacles evolving free from gastropod predation retained primitive skeletal features. This supports the interpretation that predation by drilling gastropods has been an important selective force behind the widespread, parallel evolutionary tendency towards plate reduction in the Balanomorpha; a trend recognized by Darwin (1854), but for which he offered no explanation.

In this paper, we present detailed quantitative studies of evolutionary changes over all or part of the stratigraphic ranges of five fossil radiolarian species from Pacific deep-sea sediment cores. Each of these species shows some variation of a distinctive evolutionary pattern: increase in size of measured morphologic characters, preceded and/or followed by an interval during which little or no significant change occurred.

One of the species studied (Eucyrtidium matuyamai) was allopatrically differentiated from another (Eucyrtidium calvertense). The others (Calocycletta caepa, Pterocanium prismatium and Pseudocubus vema) underwent phyletic change within a single lineage. Those species undergoing phyletic change towards larger size maintained almost constant variability of shell size over long periods of time, including periods of both rapid and extremely slow evolution. This constancy of variability suggests that diminution of selection against larger size may have acted as a stimulus to size increase. In contrast, E. calvertense decreased in variability during evolution towards smaller shell size. We believe this decrease may be interpreted as the result of two factors: (1) strong selection against larger size apparently exerted on this species by its direct descendant, E. matuyamai, during the neosympatric phase of speciation and (2) continuation of previous selection against very much smaller size.

An intertidal sample of 118 to 155 Terebratalia transversa on Saltspring Island, B.C., was censused in 1974, 1975, and 1976. Growth in length was a maximum of 7.8 mm per year and declined with size (age). As in fossil articulates, mortality rate was independent of age (size). Recruitment was patchy in time and space, was multi-annual or continuous, and was concentrated near conspecific adults. There were as many as 800 individuals/m2. None moved or changed orientation. Morphologic variation of Terebratalia valves is not directly controlled by the intensity of waves or currents.

Paleontological survivorship curves based on size-frequency data are subject to ambiguous interpretation because two critical assumptions (direct relationship of size and age, constant population structure) are difficult to justify.

We investigated the hydrodynamic properties of cephalopod shell sculpture in two ways: 1) flow visualization experiments with sculptured shells; and 2) application of drag coefficient data for simple geometric bodies to cephalopod shells. Results of this work suggest:

1) the hydrodynamic effect of shell sculpture depends primarily on the size of the sculptural elements relative to the size of the shell and on the positions of sculpture elements on the shell and relative to each other.

2) sculpture is detrimental to swimming (reduces hydrodynamic efficiency) if it exceeds the height of the lower part of the shell's boundary layer.

3) sculpture is advantageous to swimming (increases efficiency) if it remains immersed in the boundary layer and induces premature conversion to turbulent boundary layer flow. To be hydrodynamically optimal, small shells (diam ≈ 10 cm) must have rough (sculptured) surfaces, whereas large shells (diam ≈ 100 cm) require smooth surfaces. Thus, in order to maintain maximum efficiency throughout life, the ontogeny of small individuals, or species, should be characterized by progressive roughening of the shell, while large forms should become increasingly smooth. Such allometries are observed among many ammonoids.

4) sculpture always has an effect on the flow around a cephalopod shell. In some species this effect was probably negligible, while in others, those with compressed shells especially, it was probably of major importance. In these species, sculpture appears to have functioned primarily to increase swimming ability.

Medullosa stands apart from most Paleozoic seed plants in its combination of large leaf area, complex vascular structure, and extremely large water-conducting cells. To investigate the hydraulic consequences of these anatomical features and to compare them with other seed plants, we have adapted a model of water transport in xylem cells that accounts for resistance to flow from the lumen, pits, and pit membranes, and that can be used to compare extinct and extant plants in a quantitative way. Application of this model to Medullosa, the Paleozoic coniferophyte Cordaites, and the extant conifer Pinus shows that medullosan tracheids had the capacity to transport water at volume flow rates more comparable to those of angiosperm vessels than to those characteristic of ancient and modern coniferophyte tracheids. Tracheid structure in Medullosa, including the large pit membrane area per tracheid and the high ratio of tracheid diameter to wall thickness, suggests that its xylem cells operated at significant risk of embolism and implosion, making this plant unlikely to survive significant water stress These features further suggest that tracheids could not have furnished significant structural support, requiring either that other tissues supported these plants or that at least some medullosans were vines. In combination with high tracheid conductivity, distinctive anatomical characters of Medullosa such as the anomalous growth of vascular cambium and the large number of leaf traces that enter each petiole base suggest vascular adaptations to meet the evapotranspiration demands of its large leaves. The evolution of highly efficient conducting cells dictates a need to supply structural support via other tissues, both in tracheid-based stem seed plants and in vessel-bearing angiosperms.

The pattern observed in nature has often served as a compelling motivation for seeking the underlying processes which are assumed to control the pattern. Spatial and temporal patterns of species diversity are the most commonly observed and described from the study of recent and paleocommunities. These patterns include those found within a local or discrete region such as succession and changes along an environmental gradient and interregional or global patterns such as latitudinal diversity gradients and comparisons of the deep-sea and the continental shelf faunas. Many hypotheses have been advanced to explain these patterns in terms of differences in the biology of species adapted to different or changing environments. However, a more simple explanation is possible: that these patterns do not result from any specific biologic processes.

We examine this possibility and show that succession can result from probabilistic immigration and local extinction of species, diversity gradients can result from probabilistic disturbance in naturally patchy environments, and that latitudinal gradients can be simply a function of the shape of the earth. As long as such diversity patterns can be explained independent of any specific biologic processes, they offer no test of the importance of these processes, either ecologically or evolutionarily.

The traces of macroboring organisms are known throughout the Phanerozoic, with diversification and exploitation of the macroboring niche paralleling variations in the development of skeletal metazoa. The oldest macroboring biota is an abundant yet low diversity fauna in hardgrounds and reefs of Lower Cambrian age. Following the extinction of archaeocyathids at the end of the Lower Cambrian (and thus the demise of skeletal reefs until the Middle Ordovician), boring organisms appear to be restricted to submarine hardgrounds. With the development of skeletal reefs in the Middle Ordovician the macroboring fauna shows a rapid speciation and a dramatic increase in diversity. This same pattern occurs again in the Devonian. This record appears to represent refuge of the fauna in low stress, hardground environments when skeletal reefs were not present and radiation in the high stress environment of the reef when large skeletal metazoa were abundant and diverse.

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.