The living members of 113 families of bivalves and gastropods of the Californian Province include 698 species living at shelf depths, of which 538 or 77% are known as Pleistocene fossils from the same region; another 113 fossil species are extralimital, and 98 are extinct. Living species not found as fossils are chiefly rare today, and/or minute, fragile, and/or from deeper shelf habitats. Sampling of the Pleistocene record has been biased towards shallow-water assemblages. Fragile and minute forms are probably underrepresented in the record. Rare forms, however, are still appearing as new studies are conducted, and many rare species are yet to be discovered. At least 85% of durably skeletonized living species may have been captured in the record. It is probable that most durably skeletonized invertebrate species were represented in lithostratigraphic units throughout the Phanerozoic, but that this record is lost owing to erosion, burial, and destruction of skeletons in situ. The bulk of the marine invertebrate fossil record does not represent a series of unusual skeletal accumulations, but rather the preserved remnants of an excellent original record formed through ordinary though episodic processes.

Pteroscaphites are small, rare species of scaphitid ammonites from the Upper Cretaceous (Turonian-Santonian) of the Western Interior of North America. Their evolution appears to parallel that of the larger species of Scaphites or Clioscaphites, with which they co-occur. To investigate this evolutionary pattern, I constructed a phylogeny of all these species based on their distribution of shared derived characters including ornamentation, ammonitella size, whorl shape, umbilical diameter, sutures, number of whorls, adult size, shape of the adult body chamber, and apertural modifications. This analysis revealed numerous instances of congruence in the preadult morphology of the pteroscaphites and that of the co-occurring scaphites or clioscaphites. However, there is a marked divergence at maturity. Adult pteroscaphites are uniformly small (2.5 postembryonic whorls in their phragmocone) and develop apertural projections. The scaphites and clioscaphites exhibit a number of morphological changes at approximately the same whorl number, but secrete as many as two more whorls in their phragmocone before forming a mature body chamber with a relatively unmodified aperture. Both groups display sexual dimorphism. The process of progenesis may explain this conflicting pattern of congruence prior to maturity and divergence at maturity. Adult pteroscaphites are not mature replicas of the juvenile shells of the larger scaphites or clioscaphites, because maturation produces its own set of morphological modifications. Nevertheless, such features as the apertural projections on adult pteroscaphites may be interpreted as extrapolations of juvenile patterns of growth into maturity. The repeated associations of pteroscaphites with scaphites or clioscaphites in the Upper Cretaceous suggest that progenesis was iterative and may have represented an optional developmental pathway common to all of these species.

Differences in the larval ecology of Ordovician trilobites directly influenced the outcome of the Ashgill extinction (latest Ordovician) and indirectly governed the pattern of evolution in post-Ordovician trilobites. Larval ecology also affected survivorship patterns within the Ordovician, particularly between the Llandeilo and the Caradoc stages. All taxa with pelagic adults became extinct by the end of the Ordovician. Similarly, trilobites with entirely planktonic larvae had all but disappeared by the end of the Ordovician. Although suffering a significant loss in diversity, taxa with benthic larvae provided the ancestral stock for the majority of post-Ordovician lineages. Trilobites with a two-stage protaspid period (single planktonic followed by benthic larvae) suffered least during the Ashgill extinction, giving rise to approximately 23% of the new genera appearing in the early Silurian.

Patterns of extinction/survivorship among trilobite taxa with different developmental strategies indicate that the so-called Ashgill extinction was most likely the result of a composite phenomenon, including environmental perturbations, ecosystem breakdown and biogeographic restriction, and was not the consequence of a single catastrophic event (e.g., bolide collision). Indeed, our results are consistent with extinction models which invoke periodic global cooling and sea level regression associated with glaciation. Taxa with planktonic larvae first suffered a marked decline in diversity between the Llandeilo and the Caradoc (end of Middle Ordovician), coincident with the onset of the Ordovician-Silurian glaciation. During this time interval, the paleogeographic ranges of most surviving genera with solely planktonic larvae were severely constricted to lower latitude paleobiogeographic provinces. During the Ashgill (Rawtheyan to Hirnantian), climatic and sea level fluctuations were most extreme because of continued and more extensive glaciation. At this time, when extinctions were particularly severe, there was also a significant effect upon the survivorship of taxa with benthic larvae.

Our results indicate that the proportion of the life cycle spent in the water column, dependent upon a planktonic food source, was a critical factor in the survivorship of trilobite taxa during Middle to Late Ordovician time. Consequently, differences in life history patterns predisposed individual taxa to survival or extinction.

A general model of species dynamics must incorporate the effects of species number on the processes of speciation and extinction. Previous models make specific assumptions about these effects, but do not consider the effects of dynamics of lower level entities on speciation and extinction rates. A hierarchical model is developed which explicitly describes the effects of energy use by species on speciation and extinction rates. The effects of energy use are represented by parameters that characterize the average effects of energy use by each species in the biota on speciation and extinction rates. The dynamics of the model describe a sigmoidal increase in species number over time, as does the logistic model of species dynamics. However, the mechanisms of those dynamics are assumed to be different in the two models. Empirical analysis of a data set on the diversification of fossil Miocene horses suggests that the logistic and hierarchical models have similar descriptive power. The hierarchical model incorporates insights from recent considerations of the nature of a hierarchical theory of biology. Further progress in developing such a theory will depend on the success with which relationships among levels in the biological hierarchy are able to be defined.

A moving reference frame is introduced for the analysis of accretionary shell growth. Simple principles of motion and stepwise growth define the model. At each growth step, the aperture migrates from its present position to a new position, according to locally defined rules. The aperture becomes the focus of the analysis, mathematically and conceptually, in conformity with biological reality. Kinematic principles provide the analytical framework for describing the aperture's trajectory (kinematics is the study of motion). The aperture “translates,” “rotates,” and “dilates.” The model offers exceptional flexibility in the analysis of accretionary growth forms and is particularly well-suited to analysis of conical and loosely coiled shell geometries. Computer simulations illustrate the principles of a moving reference model. The inverse problem of finding the aperture motions from actual shell data is rigorously specified, for both planispiralled and helicospiralled shell forms.

A marked retardation of dental ontogeny characterizes the family Elephantidae. As a consequence of this retardation, elephant teeth are subject to the forces of mastication, eruption, and progression while still in a developing and pliant stage. As specimens described here illustrate, the mechanical forces are often sufficient to deform the gross morphology of dentitions. Morphological variation in elephant teeth can be regarded as “fabricational noise”—revealing information about the dynamic context in which the teeth develop. Accordingly, dental variation is less species-specific in elephants than in other mammals. The fossil record may comprise fewer species of elephants than is generally believed, and trends inferred to reflect rapid evolution within this family may in fact reflect phenotypic plasticity.

Application of multivariate statistical techniques, especially correspondence analysis, results in the recognition of four major communities for the Creede plant assemblages: fir-spruce forest, fir-pine forest, pine-juniper forest or woodland, and mountain mahogany chaparral. Physiognomy of the Creede assemblages indicates a mean annual temperature of <2.5°C. Paleoaltitudinal estimates based on this temperature estimate are inconsistent with physiographic and structural data of post-depositional uplift, unless the climatic effects of a closed basin are factored in; these effects imply that the Creede paleotemperature (and hence paleoaltitude) is applicable to the caldera rim. A variety of data indicate that summers were dry and most precipitation occurred as snow. The stratigraphic sequence of assemblages indicates that significant precipitational change occurred during deposition of the Creede plant-bearing beds. Chamaebatiaria, a shrub that now inhabits dry, open environments, had an ancestral taxon characteristic of the Creede fir-forest communities and strongly indicates a major shift in habitat for this lineage during the Neogene. A second genus, Luetkea, which is now herbaceous, is represented by a probable woody ancestral taxon that was common in the Creede forest communities. Consideration of the adaptive and morphologic histories of the Creede lineages suggests that physiologic adaptation may precede morphologic change. The Creede forest communities have no modern homologues.