The origin and diversification of terrestrial communities has long been associated with the Devonian. The earliest known substantial assemblage of land plants and animals preserved in or close to their original habitats is still that of the Rhynie Chert peat bog, from Scotland, which is of Siegenian age.

Two rival models of evolution in multispecies systems are tested against empirical species-level data. The two models are the Red Queen model of Van Valen as reformulated by Stenseth and Maynard Smith, which assumes that evolution is driven principally by biotic interactions, and the Stationary model of Stenseth and Maynard Smith, which assumes that evolution is propelled primarily by abiotic factors and will cease in the absence of changes in abiotic parameters. Testing refers to the models' predictions regarding the behavior of extinction and origination rates, and assumptions regarding equilibrium diversity and a constant effective environment. The data set includes the dates of origination and extinction for all coccolith, planktic foraminifer, and radiolarian species recorded in the Oligocene through Holocene, and all planktic diatom and silicoflagellate and ebridian species recorded in the Middle Miocene through Holocene in 111 DSDP sites of the low- to mid-latitude Pacific Ocean.

The condition of stable specific age distribution over geologic time is met, which allows one to perform survivorship analysis on extinction rates. The best fit survivorship curve is a decreasing function of age for both coccolith and foraminifer species, and an increasing function of age for radiolarian species. Neither model predicts age dependence of the probability of extinction. The small disparity between these curves and age-independent curves for each group indicates, however, that an age-independent interpretation of extinction probability is a reasonable first approximation. Rates of origination are analyzed in terms of species accretion, introduced to represent the cumulative origination of species within a higher taxon as a function of the age or duration of the community. Accretion analysis indicates that the probability of accretion is both diversity-dependent and absolute time-dependent.

The assumption of a constant effective environment is tested by polycohort analysis and nonparametric logistic regression analysis of true species cohorts. Both techniques indicate considerable variation in extinction probability over geologic time. When the predictions of the two evolutionary models are adjusted to take this variation into account, the results of both survivorship and accretion analysis seem to conform more closely to the predictions of the Red Queen than to the Stationary model. However, as the speed with which the effective environment changes is increased relative to speciation-extinction rates, it becomes increasingly difficult to differentiate patterns predicted by the two models. The assumption of equilibrium diversity can be neither corroborated nor rejected, since the species-level data are compatible with both an equilibrium and a nonequilibrium view of diversity behavior. Reservations concerning the basic assumptions of both models indicate an ultimate test requires that both models be reformulated to make precise and distinctive predictions under a varying effective environment.

Available data on the stratigraphic ranges of latest Silurian and Devonian vascular plant macro-fossils (sporophytes) and spores provide insights into the tempo and mode of early tracheophyte evolution. Patterns of diversification, origination, and extinction conform in general to the predictions of Sepkoski's kinetic model of diversification. Rates of generic origination and extinction vary not only through time but also between organ systems for a single time interval. This fact, coupled with data on longevity and turnover and comparative morphological observations, can be used to document mosaic evolution in early vascular plant history. Mosaic evolution is an important theme in plant evolution; indeed, what we recognize as macroevolutionary events often correlate with brief periods of pronounced mosaicism. Such evolutionary patterns reflect the developmental biology of tracheophytes in which individual organs often have life spans that are considerably shorter than the life of the whole plant. Under these conditions, individual organs or organ systems can respond to different sets of evolutionary pressures.

The major period of early vascular plant diversification occurred during the late Early and early Middle Devonian Period, 30 Myr or more after the origin of the group. Such lags in diversification are not uncommon in the fossil record. Sometimes they reflect extrinsic controls on diversification, but in other cases they appear to be a consequence of intrinsic rates of origination and extinction.

Faunal stasis among Indo-Pacific reef-building corals is explained as the result of chronic environmental disruptions preventing evolutionary processes from approaching completion since the Late Pliocene. The model assumes shallow reefal habitats (<20 m) on the continental shelves are major sites of scleractinian evolution and explores ecological and evolutionary consequences of high-frequency sea-level fluctuations and their associated transgression-regression cycles. Because single generations, dominated by a few large clonal genotypes, may persist indefinitely, local populations may not have experienced enough generations to approach evolutionary equilibrium with their environments during the estimated average duration (≈3200 yr) of existence of shallow habitats. Persistent consequences of chronic evolutionary disturbance may be the extensive intraspecific variation so characteristic of the dominant genera of shallow Indo-Pacific corals and the apparent paucity of recently evolved endemic species. The same disturbances may have accelerated speciation rates among reefal organisms with much shorter generation times.

Surprisingly, there is a relationship between rates of sediment accumulation and the time spans for which they have been calculated. This relationship can be used to estimate expected rates for specific sedimentary environments and time spans. The most probable completeness of a given sedimentary section at a given short time span can be calculated by the ratio of the measured long-term rate of sediment accumulation to the expected short-term rate. Although the measured time span is usually based on radiometric and paleomagnetic data, the cumulative time of formation estimated from fossil soils in a sequence may also be used to calculate rates and may be useful in comparing the completeness and rate of accumulation of different sequences. By both kinds of estimates, terrestrial sedimentary successions are disappointingly incomplete. Some reasons for incompleteness are illustrated with a simple model of episodic flooding, exceeding a threshold for destruction and sedimentation over a particular kind of vegetation, and thus initiating a new cycle of soil formation. In such a model, rock record is lost to erosion during cutting and filling cycles, to overprinting of weakly developed soils by later, better-developed soils, and to continued development, near steady state, of the soils preserved.

Because fossil soils are evidence of ancient environments and ecosystems independent of the fossil record, they may provide evidence of expected kinds of fossils, such as silica phytoliths, calcareous phytoliths, pollen, leaves, fruits, seeds, charcoal, land snails, coprolites, and bones. The degree to which the kinds of fossils actually found fail to meet these expectations is a crude measure of the completeness of representation of a former ecosystem in the fossil record. Some of the discrepancy between expected and actual occurrence of fossils can be related to the original Eh and pH of a fossil soil, as approximated by the oxidation state of iron in its minerals (for Eh) and by carbonate or zeolite content (for pH). Different kinds of fossils can be envisaged as having a characteristic Eh-pH stability field within which they can be expected to have been preserved if originally present. Even under ideal conditions of preservation, it takes some time for fossils to accumulate in soils to levels at which representative collections can be made. Estimates of this temporal control on preservation can be gained by comparing fossil occurrences with the degree of development of fossil soils. Neither these chemical nor temporal factors account fully for the degree of incompleteness observed because original abundance, trampling, predation and many other factors are also important determinants of fossil occurrence.

These considerations can be used as guidelines for choosing stratigraphic sections appropriate for particular paleobiological and geological problems. For example, a study of speciation of terrestrial vertebrates would best be in a sequence of weakly developed, calcareous fossil soils (Entisols and Inceptisols), of near-uniform texture and yellow to brown color, formed under an extraordinarily high long-term rate of sediment accumulation. On the other hand, a study of coevolution of vertebrates and plants would best be based on a sequence of weakly to moderately developed, calcareous fossil soils of predominantly drab (gray, green, and blue) color, with interbedded carbonaceous shales.

A computer model is presented which is capable of calculating both the photosynthetic efficiency (I) of any specified plant shape and the stress related to the total moment arm (M) imposed on vertical branching patterns. Computer simulations indicate that a flattened plant thallus and an erect branching growth habit are two plant shapes capable of optimizing photosynthetic efficiency during indeterminate growth. These two morphologies have geometric analogues in the dorsiventral thalli of some bryophytes and in the vertical axes of mosses and tracheophytes, respectively.

Extension of the model to complex, three-dimensional branching patterns indicates that I and I/M are maximized when branching is overtopped (treelike, with lateral branches on a main axis) and when lateral branching systems are planated (frondlike). Geometric alterations of branching patterns that result in optimization of I and I/M can be simulated by computer and are shown to be similar to morphologic alterations attending the early evolution of vascular land plants. It is suggested that a number of major evolutionary trends seen in Upper Silurian to Upper Devonian times can be expressed in terms of optimizing the display of photosynthetic tissues (I) or the balance between photosynthetic efficiency and incurred moment arms (I/M).

Three features that provide information on the paleobiologic significance of fossulae in North American Late Ordovician solitary rugose corals are (1) irregularities in shape and position of the outer wall, (2) discontiguous septal growth lamellae, and (3) foreign objects incorporated into interseptal chambers within the corallum. Outer wall irregularities indicate that portions of a polyp could detach from the calice and could contract and expand radially by a significant amount and for prolonged periods of time during ontogeny, especially in the vicinity of the alar and cardinal fossulae. Discontiguous lamellae indicate that the polyp could detach from septa and contract laterally, especially in the cardinal fossula. This began with the onset of maturity and probably reflects reproductive activity. Ostracodes are the most common foreign objects in these coralla and are usually situated within or near the cardinal and alar fossulae. In the most likely hypothesis accounting for their presence, a live ostracode entered the calice when one side of the polyp temporarily detached from the corallum and contracted radially. It became trapped upon expansion and reattachment of the soft parts. The polyp moved upward in its corallum by detachment and uplift of the aboral surface. In a less likely hypothesis, the ostracode was captured by the coral for food and came to rest on the floor of the central cavity. It was incorporated into an interseptal chamber when the polyp moved upward in its corallum by atrophy of the aboral surface and formation of a new base above the object. In either hypothesis, portions of the polyp in the cardinal and alar fossulae probably functioned throughout ontogeny for water circulation in the central cavity, and for the intake of food and/or ejection of undigested material through the mouth. Taxonomic, stratigraphic, and paleobiogeographic variability in frequencies of the three features may indicate differences in the necessity and/or ability of polyps to perform these functions involving the fossulae. This could be a reflection of environmental and/or genetic factors.