Adaptation has been defined and recognized by two different criteria: historical genesis (features built by natural selection for their present role) and current utility (features now enhancing fitness no matter how they arose). Biologists have often failed to recognize the potential confusion between these different definitions because we have tended to view natural selection as so dominant among evolutionary mechanisms that historical process and current product become one. Yet if many features of organisms are non-adapted, but available for useful cooptation in descendants, then an important concept has no name in our lexicon (and unnamed ideas generally remain unconsidered): features that now enhance fitness but were not built by natural selection for their current role. We propose that such features be called exaptations and that adaptation be restricted, as Darwin suggested, to features built by selection for their current role. We present several examples of exaptation, indicating where a failure to conceptualize such an idea limited the range of hypotheses previously available. We explore several consequences of exaptation and propose a terminological solution to the problem of preadaptation.

Barstovian (medial Miocene) mammalian faunas from the Texas Gulf Coastal Plain contained four apparently sympatric species of rhinoceroses: the common forms Aphelops megalodus and Teleoceras medicornutus, a dwarf Teleoceras, and a dwarf Peraceras. Previous work has suggested positive allometry in tooth area with respect to body size in several groups of mammals, i.e., larger mammals have relatively more tooth area. However, dwarfing lineages were shown to have relatively more tooth area for their body size. Our data show no significant allometry in post-canine tooth area of either artiodactyls or ceratomorphs. Similarly, dwarf rhinoceroses and hippopotami show no more tooth area than would be predicted for their size. Limbs are proportionately longer and more robust in larger living ceratomorphs (rhinos and tapirs) than predicted by previous authors. Limb proportions of both dwarf rhinoceroses and dwarf hippopotami are even more robust than in their living relatives.

The high rhinoceros diversity reflects the overall high diversity of Barstovian faunas from the Texas Gulf Coastal Plain. The first appearance of several High Plains mammals in these faunas indicates “ecotone”-like conditions as faunal composition changed. Study of living continental dwarfs shows that there is commonly an ecological separation between browsing forest dwarfs and their larger forebears, which are frequently savannah grazers. This suggests that the dwarf rhinoceroses might have been forest browsers which were sympatric with the larger grazing rhinos of the High Plains during the Barstovian invasion. The continental dwarf model also suggests that insular dwarfism may be explained by the browsing food resources that predominate on islands.

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.

Species diversity is a product of both biological interactions and physical-geographic factors. A model which eliminates biological interactions as a determinant of diversity can be used to assess the potential impact of physical and geographic factors operating alone on temporal trends in diversity (succession) and on spatial differences along latitudinal or environmental gradients. If a biologically neutral model can generate diversity patterns similar to those we observe, it can be argued that traditional hypotheses which depend on biological interactions are not the most parsimonious explanations for diversity patterns. If the neutral model approach supports a spectrum of possible hypotheses, perhaps the entire concept of species diversity can be discarded as a useful measure of community structure.

An earlier analysis of this type supported the notion that diversity patterns can be adequately explained by physical disturbance, global geography gradients, and stochastic immigration and extinction events. Our neutral model analysis does not affirm this view. We argue that the coarse level at which simple colonization models operate does not permit strong conclusions regarding the underlying mechanisms. Detailed community analyses may provide the best evidence for the nature of structuring processes. Based on the evidence available in the literature, species diversity is likely to be generated by a small number of biological processes interacting with physical-geographic gradients.

Phyletic speciation arising from a shift between adaptive peaks in the selective surface is modeled. At a locus initially monomorphic, a new allele is introduced that triggers phenotypic evolution by making a new adaptive peak available. A phenotypic model is used to describe the evolution of a quantitative character that is important for fitness but not directly affected by the new allele. Recursion equations are derived for both the allele frequency and the mean of the quantitative character. It is shown that the dynamics of the quantitative trait fit a “punctuated equilibrium” pattern of evolution. Phyletic speciation is seen to occur over a time span so short as to make observation of the transition in the fossil record unlikely.

Substrata for numerous epizoans in the Silurian Waldron Shale were provided by the toppled and overturned calyces of camerate crinoids. These were inhabited by a skeletonized fauna consisting of at least 25 species; including several bryozoans, tabulates, inarticulate brachiopods, worms, and echinoderms.

Many of the encrusting bryozoans exhibit skeletal overgrowths which appear to reflect competitive interactions for space. Coexistence of certain of the bryozoans involved in such interactions is suggested by skeletal features such as inflated growth forms and upturned margins at junctions between colonies (both representing attempts to avoid being overgrown). In addition, the occurrence of overgrowth reversals within individual encounters indicates contemporaneity of that pair. Species may be ranked according to their relative overgrowth success; however, this ranking is far from rigid as lower ranked species may on occasion overgrow more highly ranked species. Such nontransitive patterns appear similar to those described from Recent cryptic bryozoan faunas.