We believe that punctuational change dominates the history of life: evolution is concentrated in very rapid events of speciation (geologically instantaneous, even if tolerably continuous in ecological time). Most species, during their geological history, either do not change in any appreciable way, or else they fluctuate mildly in morphology, with no apparent direction. Phyletic gradualism is very rare and too slow, in any case, to produce the major events of evolution. Evolutionary trends are not the product of slow, directional transformation within lineages; they represent the differential success of certain species within a clade—speciation may be random with respect to the direction of a trend (Wright's rule).

As an a priori bias, phyletic gradualism has precluded any fair assessment of evolutionary tempos and modes. It could not be refuted by empirical catalogues constructed in its light because it excluded contrary information as the artificial result of an imperfect fossil record. With the model of punctuated equilibria, an unbiased distribution of evolutionary tempos can be established by treating stasis as data and by recording the pattern of change for all species in an assemblage. This distribution of tempos can lead to strong inferences about modes. If, as we predict, the punctuational tempo is prevalent, then speciation—not phyletic evolution—must be the dominant mode of evolution.

We argue that virtually none of the examples brought forward to refute our model can stand as support for phyletic gradualism; many are so weak and ambiguous that they only reflect the persistent bias for gradualism still deeply embedded in paleontological thought. Of the few stronger cases, we concentrate on Gingerich's data for Hyopsodus and argue that it provides an excellent example of species selection under our model. We then review the data of several studies that have supported our model since we published it five years ago. The record of human evolution seems to provide a particularly good example: no gradualism has been detected within any hominid taxon, and many are long-ranging; the trend to larger brains arises from differential success of essentially static taxa. The data of molecular genetics support our assumption that large genetic changes often accompany the process of speciation.

Phyletic gradualism was an a priori assertion from the start—it was never “seen” in the rocks; it expressed the cultural and political biases of 19th century liberalism. Huxley advised Darwin to eschew it as an “unnecessary difficulty.” We think that it has now become an empirical fallacy. A punctuational view of change may have wide validity at all levels of evolutionary processes. At the very least, it deserves consideration as an alternate way of interpreting the history of life.

The distribution of numbers of species and the median number of species from 386 selected fossil communities are tabulated for high stress, variable nearshore, and open marine environments during the Lower, Middle, and Upper Paleozoic, the Mesozoic and the Cenozoic. The number of species always increases from high stress to variable nearshore to open marine environments. Within-habitat variation in number of species is small for long intervals of the Phanerozoic. The median number of species in communities from high stress environments remains fixed at about 8 from the Cambrian to the Pleistocene. In open marine environments, the median is near 30 for the Middle and Upper Paleozoic and almost the same for the Mesozoic. Increases of 50% in median number of species between the Lower and Middle Paleozoic and 2 times between the Mesozoic and Cenozoic occur in open marine environments with parallel, but less pronounced, increases in variable nearshore environments. Conditions controlling overall within-habitat species richness changed at those times. These changes do not correlate directly with evolution of new major taxa, change in physical conditions, predation, space availability or oxygen supply. They may be related to changes in resource availability influenced by factors such as the developing terrestrial flora, to lag-time inherent in the evolutionary process of diversification, or to as yet undetermined factors. Although provinciality determines total species richness for the biosphere, the within-habitat data suggest that the number of marine invertebrate species in the world has increased since the Middle Paleozoic, contrary to Raup's (1976b) contention, but possibly only by about 4 times, not the order of magnitude or more suggested by Valentine (1970).

The fossil record of echinoids is poor because soon after death they break apart into isolated plates. In the present seas regular echinoid species outnumber irregular species; whereas, in the Tertiary only 20% of the known echinoid species are regular. This suggests that regular echinoids are less likely to be preserved than irregular echinoids. The tests of regular echinoids are exposed to scavengers and currents upon their death, but irregular echinoids generally live buried in the sediment and are protected from these destructive forces. Furthermore, the tests of the regular echinoids lack the calcareous supports found in some irregular echinoids. The gut is not filled with sediment and its apical system is generally larger and more fragile. Finally, many regular echinoids live in environments less likely to be preserved in the sedimentary record.

Although the irregular echinoid is more likely to be fossilized, its record is poor during some periods in the past. Although 1,014 irregular echinoid species are known from the Eocene, only 83 species are known from the Paleocene and only 343 are known from the Oligocene. Is this reduction because fewer species lived then or because they have not been preserved?

Euproops danae, a xiphosuran from the Mazon Creek (Braidwood) Fauna, had two sets of spines—genal and ophthalmic—extending from the posterior margin of its prosoma. In young individuals, the genal spines were longer than the ophthalmic spines. During growth, the relative length of each type of spine increased, but the rate of increase was much greater for ophthalmic spines. In order to explain these morphological and ontogenetic features, I have studied the hydrodynamic behavior of whole-animal models representing the actual morphology of E. danae, and various modifications thereof, at different body sizes. It can be argued that passive settling, while in an enrolled posture, was probably the primary defensive reaction that a swimming individual would have exhibited in response to an encounter with a potential predator. Experiments show that the array of spines on E. danae was an important control on the style of settling experienced by an enrolled individual. Models of the actual morphology settle steadily, while models with either longer or shorter prosomal spines tend to undergo lateral oscillations induced by turbulence in the wake. Steady fall would have rendered the horseshoe crab less perceptible to either the visual or lateral line systems of contemporary aquatic vertebrates, and thus a morphology capable of producing it would have been an important adaptation for reducing the risk of predation. This minimization of oscillatory movement would actually obtain for a variety of conceivable spine morphologies, but actual morphologies represent those ‘solutions’ which simultaneously optimize other aspects of spine function, such as mechanical protection. This optimum design changes during ontogeny because settling dynamics scale non-linearly with size. This work has both specific applications to the interpretation of similar structures in other arthropods and more general implications for the study of evolutionary functional morphology.

Five generalizations derived from the shell form of prosobranch gastropods are developed. (1) A univalve of more than one volution whose aperture lies in a plane that passes through the axis of coiling does not live with the aperture parallel to the substrate. (2) A univalve of more than one volution whose aperture lies in a plane that is tangential to the body whorl does live with the plane of the aperture parallel to the substrate. (3) Gastropods with tangential apertures, when extended, support the shell so that the center of mass of the shell and its contents is over the midline of the cephalopedal mass; this balancing of the shell may be accomplished either by regulatory detorsion, by inclination or by a combination thereof, to keep the center of gravity of the shell as low as possible. (4) Angulations or re-entrants in the gastropod aperture are usually indicative of inhalent or exhalent areas; inhalent areas are directed as far anteriorly as possible. (5) Gastropods having elongated apertures possess only a single gill and develop a water current through the mantle cavity from anterior to posterior along the long axis of the aperture; this axis is subparallel to the anterior-posterior axis of the foot.

These generalizations are then used as the basis for some deductive interpretations of behavioral modes of Paleozoic Gastropoda.

Silurian crinoids reached maximum abundance on shallow shelf bottoms of skeletal-rich carbonate. In-place crinoid preservation in the Wenlock Limestone of Dudley, England, provides a community model for these occurrences, showing a pattern among crinoids of high taxonomic diversity, conspecific clustering, relatively robust morphology, and numerical dominance of other invertebrates. Densities of about 40 crinoid individuals per m2 occupied a feeding level between 10 cm and 1 m above the bottom and may have effected trophic control over other fauna. Beneath the crinoids lived a second level of bryozoa, a third level of tabulates, and a sparse fauna of brachiopods and other solitary organisms. Crinoids also exercised significant biologic control of substrate. In contrast to the Dudley assemblage, Silurian communities of terrigenous, soft-bottom environments contained crinoids of low abundance, low taxonomic diversity, and small size. In the terrigenous communities, crinoids appear to have had no major effect on either trophic structure or substrate. High sedimentation rates limited the role of crinoids in such communities, and the dominant organisms were deposit feeders and solitary, low-level suspension feeders.

A major aim of the analysis of faunal remains from archaeological sites is the reconstruction of the palaeoecology of early man. Since taphonomic work must precede palaeoecological work, theories concerning the palaeoecology of early man would benefit from firmly based taphonomic studies of the archaeological deposits. One of the numerous taphonomic factors to be considered is the bone fragmentation produced by early man.

The degree of bone fragmentation in a deposit uninfluenced by human factors is an expression of the resistance of the various bones to mechanical and chemical decomposition before and during burial. Thus, the degree of fragmentation where human factors are predominant should provide an illustration of man as a taphonomic factor. In both cases the degree of fragmentation may be used as an indicator of the degree of taphonomic overprint.

Examination of bone material from four West European Mesolithic sites (all bog deposits), Star Carr 7200 ± 120 B.C., Kongemosen 6600 ± 100 B.C., Præstelyngen 3200 ± 100 B.C., and Muldbjerg I 2900 ± 80 B.C., reveals differences in marrow fracturing techniques. Furthermore, the different techniques result in different numbers of fragments for the same type of bone. Comparison of faunas from different sites based on the number of bone fragments must therefore be preceded by analysis of the marrow fracturing techniques used at the various sites. The number of fragments per estimated minimum number of individuals should indicate the degree of taphonomic loss.

The various types of marrow fracturing found seem to belong to cultures at different levels of technical development. Thus, it may be possible within a limited area such as northern Europe to establish a chronology using marrow fracturing type as one of the ecostratigraphic tools.