Fourier analysis of percent extinction data for genera of fossil marine animals during the past 260 million years indicates the existence of a 26 m.y. extinction cycle. The first three harmonics, which account for 51.7 percent of the sum of squares, show the major extinction trends in the late Paleozoic, Mesozoic and Cenozoic. The first ten harmonics, which account for 89.5 percent of the sum of squares, are aligned with eight extinction peaks. The tenth harmonic, with a period of 26 million years, has an amplitude of 6.5 percent and accounts for 12.7 percent of the sum of squares.

Several different methods are used to test the validity of the apparent 26 m.y. extinction cycle. Analysis of variance indicated that the 26 m.y. extinction cycle is statistically significant at the 95 percent level. When the 260 m.y. sample interval is divided into equal 130 m.y. segments, the phases and amplitudes for the 26 m.y. cycle were almost equal within the two independent segments. When the 260 m.y. interval is divided into 10 shorter time segments of different lengths, the 26 m.y. cycle is represented in each time segment.

A series of experiments is run to test the influence of the Late Permian and Cretaceous peaks on the observed 26 m.y. cycle. When all peaks are smoothed out except for the two major peaks, a pseudo-cycle with a period of 26 m.y. is generated that accounts for 3.0 percent of the sum of squares. When the Permian and Cretaceous peaks are reduced to about half their observed height, the sum of squares accounted for by the 26 m.y. cycle is reduced from 12.7 to 11.0 percent. When the number of extinctions within each stage is used in the harmonic analysis in place of extinction percent, the 26 m.y. cycle accounts for 19.3 percent of the sum of squares.

Therefore, the evidence from harmonic analysis of fossil marine animals points toward a distinct and persistent 26 m.y. cycle in mass extinctions in the late Paleozoic, Mesozoic and Cenozoic.

We have calculated rates of evolution for 19 lineages of Neogene bivalves over time intervals ranging from 1 ma to 17 ma. Our morphometric comparisons are based on 24 variables, for which we have made more than 43,000 individual measurements normalized for shell size. We initially assessed evolutionary changes in shape for 19 early Pliocene (4 ma old) species of bivalves by comparing these forms to their closest living relatives, which in 12 cases have traditonally been assigned to the same species. To make our study unbiased and comprehensive, we included all species that met certain predetermined guidelines and that offered suitable fossil material for measurement. We compared early Pliocene and Recent populations using (1) all 24 variables treated separately, (2) multivariate distance (Mahalanobis' D), based on the full set of variables, and (3) eigenshapes for shell outlines. For these comparisons, we used as a yardstick the same measures of morphologic distance applied to pairs of geographically separated Recent populations that belong to eight of the living species to which the fossil populations were compared. As it turns out, with minor exceptions, the distribution of morphologic distances between 4 ma old and Recent populations resembled the distribution of distances between conspecific Recent populations.

We calculated net rates of evolution separating pairs of populations that belong to single lineages. For all intervals of time, the distribution of differences between population means for individual variables is remarkably similar to a comparable distribution representing the comparison of pairs of conspecific Recent populations from separate geographic regions. Because morphologic differences between populations do not vary greatly with evolutionary time, measured “rates” of evolution, on the average, decrease with interval of measurement. Because these differences resemble intraspecific variability, however, the rates do not represent significant evolution. Evolution has followed a weak zigzag course, yielding only trivial net trends.

The weak and reversible “trends” that we measured yield net rates averaging less than 10 millidarwins, which is much lower than most rates previously reported for marine invertebrates (average ~200 millidarwins for a 1 ma interval and ~60 millidarwins for a 10 ma interval). We attribute this disparity (1) to the fact that most previously published rates have been calculated when a significant amount of evolution was recognized in advance (often for a poorly documented lineage) and (2) to the fact that most measured variables have represented nothing more than some measure of body size. We conclude that shape, as opposed to size, has been highly stable in bivalve evolution over millions of years and 106–107 generations. We conclude that to characterize rates or evolution for any group of organisms, one must employ a large, unbiased sample of measurements for numerous well-documented lineages, and one must segregate data depicting shape from data depicting size.

We have produced computer simulations of multiramous graptoloids with the intention of defining the rules governing branching strategies and colony form. Close matches between such simulations and real graptolites show that complex rhabdosomes may be produced by the permutation of relatively simple sets of rules. Those designs found in nature produce an efficient and regular distribution of zooids through the area included by an essentially planar rhabdosome. Strikingly geometrical arrays of stipes, such as the Goniograptus and yin/yang patterns, closely approach paradigmatic harvesting arrays. For dichotomously branching anisograptids the evolutionary trend in reduction of “primary stipes” can be explained by the production of larger spreading colonies. Multiramous graptoloids fed during vertical movement through the water column. Changes in a single branching decision may produce considerable changes in rhabdosome morphology, but these are not necessarily of high taxonomic importance; this is proved by a specimen which is a morphological combination of two “genera.” Although primarily under genetic control, certain modifications to colony form were probably the result of inhibitory interaction between adjacent stipes.

The occurrence of zooxanthellae in Recent scleractinian corals is strongly correlated with their growth form, corallite size, and degree of morphological integration of corallites. The great majority of zooxanthellate corals are multiserial with small, highly integrated corallites, whereas most corals lacking zooxanthellae are solitary or uniserial colonial forms with large, poorly integrated corallites. Beginning in the Jurassic, fossil scleractinian faunas are morphologically similar to Recent faunas dominated by zooxanthellate species, strongly implying that most scleractinians contained zooxanthellae by that time. Evidence for Siluro–Devonian tabulates and Triassic scleractinians is equivocal but still suggests the presence of zooxanthellae in these corals. In contrast, morphological evidence suggests that rugosan corals lacked zooxanthellae.

Most populations of Recent zooxanthellate corals contribute to reef formation, but many do not. Similarly, fossil corals interpreted to contain zooxanthellae on morphological grounds did not always form reefs. Recent reef formation depends upon a host of environmental factors that have little to do with the possession of zooxanthellae per se. Coral morphology should be a better predictor of the presence of zooxanthellae in fossil corals than their association with reefs.

Astogenetic trajectories have constrained evolutionary changes in bryozoans. Rates and timing of astogenetic differentiation have been modified for characters defining the morphology of zooids, subcolonies, and colonies. This paper catalogs 46 examples of bryozoan heterochrony, representing all five skeletonized orders. Heterochrony is inferred to have been a pervasive phenomenon in the evolution of Paleozoic stenolaemates, illustrated by 40 examples, 19 of which produced paedomorphosis and 21 peramorphosis. As a consequence, a restricted range of morphologic states has reappeared repetitively as homeomorphies and evolutionary reversals. Large-scale patterns developed across both geologic time and geographic space reflect variation in heterochronic products irrespective of the developmental processes by which they were achieved. Available evidence indicates that smaller, paedomorphic, and more plastic species inhabited onshore, low-diversity areas. Nonheritable plasticity is inferred to be a correlate of early growth stages and paedomorphosis. Taller, generally peramorphic species with damped plasticity are found in higher diversity, offshore regions. Seven key innovations, which first appeared during the early diversification of bryozoan clades, are peramorphic, and recapitulation was a predominant pattern during their Ordovician radiation. Trends in later phylogeny, on the other hand, have favored paedomorphic derived morphologies, as illustrated by 19 of the 32 examples. Recurrent reverse recapitulation suggests that offshore ancestors frequently gave rise to onshore paedomorphs.

This study addresses the question of evolutionary tempo and mode in a sequence of Upper Cretaceous bivalves in the genus Pleuriocardia from the Western Interior Basin of North America. Change between species was probably phyletic (without persistence of ancestors). There is some evidence for weak gradual change within the lineage, but most important change is concentrated in short intervals of time. Detailed examination of the differences among samples reveals pronounced geographic variation, whereas temporal variation within localities is generally minor. The relatively rapid episodes of change fit the model of punctuated equilibrium, but the phyletic nature of species-level change does not.

The value of the debate about the punctuated and gradualistic models has been to force a more criticial examination of fossil sequences, but these sequences should not be forced into narrowly defined categories. A variety of evolutionary patterns may exist in the fossil record, and it is this variation in pattern that will inform us of the underlying processes.

Distributional patterns of late Osagean (Mississippian) crinoids from the east-central United States are examined using multivariate analysis of crinoid species diversity and species abundance data. We confirm previous hypotheses that three well-defined crinoid associations existed during the late Osagean. These associations were dominated, respectively, by 1) monobathrid camerates preserved in carbonate packstones; 2) poteriocrine inadunates in higher-energy siltstones and sandstones; and 3) disparid inadunates, cyathocrine inadunates, and flexibles in mudstones where neither monobathrids nor poteriocrines dominated. In conjunction with petrologic data on the enclosing sediments, the analyses suggest that these associations occurred along a spectrum of increasing current velocity at the seafloor. Camerates, poteriocrine inadunates, and flexibles are interpreted to display preferences for specific environmental conditions, whereas disparid and cyathocrine inadunates are inferred to be environmental generalists.

The different environmental distributions of the major crinoid groups are interpreted to be a function of the mode or modes of aerosol filtration feeding used by each group. This inference is possible through functional morphologic and morphometric studies of crinoid arms, because the skeletal elements of the arms, which are commonly preserved, are directly involved in feeding.

A new method of data analysis offers a potentially powerful tool for statistically evaluating hypotheses of rate in temporally-ordered evolutionary phenomena. We present a method for bootstrapping time-ordered data sets to test hypotheses of the equality of rate. This method is applicable to both nonrandom and random generative processes. The method is applied to the data of Malmgren et al. (1983) for the Globorotalia plesiotumida–G. tumida planktonic foraminiferan lineage and the data of Reyment (1982) for the benthonic foraminiferan Afrobolivina afar. G. plesiotumida is recognizable on the basis of independent data as a species distinct from G. tumida, its descendant. Evolutionary change rate during the evolution of G. tumida from G. plesiotumida is shown to be faster than rates within either species. The pattern of variation exhibited by A. afar includes a time interval of more rapid change; this more rapid change is observed post hoc. A bootstrapping model based on post hoc observations reveals the rate in this time interval to be not significantly faster than expected in such post hoc intervals.

The known Cretaceous formicoids are better interpreted from morphological evidence as forming a single subfamily, the Sphecomyrminae, and even a single genus, Sphecomyrma, rather than multiple families and genera. The females appear to have been differentiated as queen and worker castes belonging to the same colonial species instead of winged and wingless solitary females belonging to different species. The former conclusion is supported by the fact that the abdomens of workers of modern ant species and extinct Miocene ant species are smaller relative to the rest of the body than is the case for modern wingless solitary wasps. The wingless Cretaceous formicoids conform to the proportions of ant workers rather than to those of wasps (Figs. 1–2) and hence are reasonably interpreted to have lived in colonies.

The Cretaceous formicoids are nevertheless anatomically primitive with reference to modern ants and share some key traits with nonsocial aculeate wasps. They were distributed widely over Laurasia and appear to have been much less abundant than modern ants.

The morphology of the tarsi, hindlimbs, and pelves of the earliest crocodilians and their nearest relatives, Hallopus and the “sphenosuchians,” indicates that these animals had adaptations for erect posture. The widespread distribution of apparently homologous adaptations for erect gait among the archosaurs with crocodile-normal tarsi suggests that those structures are plesiomorphic for this group, which comprises the Aetosauria, “rauisuchians,” “sphenosuchians,” Hallopus, and the Crocodylia. Adaptations for erect posture are seen most clearly in the structure of the proximal tarsus (astragalus and calcaneum).

An important implication of this argument is that the most primitive crocodylomorphs, comprising the “protosuchian” crocodiles, the “sphenosuchians,” and Hallopus, had an erect stance and gait. The sprawling stance and associated gait used by modern crocodilians during swimming and upon entering the water can be viewed as secondary adaptations to an aquatic existence. The environments of deposition and faunal associations of “sphenosuchians” and “protosuchian” crocodiles are consistent with primarily terrestrial habits. Living crocodilians have two types of step cycles, sprawling and erect; the sprawling pattern is overprinted onto the inferred ancestral “high-walk,” and onto the gallop sometimes used by juvenile crocodilians.

The extent to which perceived patterns of evolution are affected by the use of single characters versus overall morphology or measured versus counted and coded characters is explored empirically, employing multiple-character data from closely spaced sequential populations of the Neogene bryozoan Metrarabdotos. In all nine species examined, the pattern of evolution in overall morphology revealed by discriminant analysis is so highly punctuated that changes within species do not vary significantly from zero. Among the 46 single characters in the nine species, a few statistically significant temporal trends do occur, as apparent departures from the static pattern in overall morphology. However, these exceptions are so poorly related to the morphology that distinguishes inferred ancestor and descendant species from each other, and are so few in number, that they can be interpreted as chance variation from a general condition of no change within species. There is no difference in behavior between measured characters and counted or coded ones, in part because the distinction between the two types of characters can be artificial in bryozoans and other modular organisms. The results suggest that interpretation of single-character changes, in isolation rather than as statistical samples of the change in overall morphology, should be made with caution.

A diversity and turnover analysis has been undertaken for a number of invertebrate groups in the Liassic of northwest Europe. There is a more or less steady rise in diversity from the early Hettangian through to the Pliensbachian, followed by a marked decline into the early Toarcian, after which it tends once more to increase. Ammonites stand out from the other invertebrates as having had an exceptionally high rate of turnover, with very short species durations.

Increase of neritic habitat area due to rise of sea level, and recolonization following the end-Triassic mass extinction event appear to be the promoters of diversity increase or radiation. Severe reductions of neritic habitat area with associated environmental deterioration, related either to episodic marine regressions or spreads of anoxic bottom waters, and bound up respectively with sea-level fall and rise, are seen as the prime factors responsible for increase of extinction rate. While the environmentally sensitive ammonites were affected by even minor regressions, the other, more eurytopic groups were evidently more resistant to these. The only event that warrants the term mass extinction, affecting nearly all the benthos and nekton but not the plankton, correlates precisely with the early Toarcian anoxic event. Several episodes can be recognized of migrations of organisms into Europe following extinctions.

The unique holotype of the Triassic reptile Sharovipteryx (initially Podopteryx) mirabilis has been reexamined and redescribed, correcting the original account. It is a small lizard-like reptile with an elongate head and densely tubular femur and tibia each longer than the (estimated) intergirdle distance. (Part of the pectoral region has been lost and the remnants remain encased in the matrix.) The matrix retains the impression of portions of the integument, including a patagium that reached from the hindlimbs to the base of the tail. Unlike the original account, the patagium did not extend to the pectoral limbs. Experiments with models indicate that Sharovipteryx could have maintained a shallow glide if the femora were held at a shallow angle to the vertebral column and the tibia and feet extended out at right angles to it, thus stretching the integument. The pectoral limbs (with or without a fringing membrane) might have produced a variable canard; alternatively vertical bending of the tail could have applied drag, each making the glide more stable.

Species duration data for living benthic foraminifera derived from an extensive literature search have been compiled and analyzed to investigate rates and patterns of species origination. The same data subjected to taxonomic standardization through examination of many specimens lodged in museum collections indicate strikingly different, and more realistic, rates and patterns.

Evolutionary generalizations based on data generated from the literature only are often unreliable and may be directly in opposition to reality. Extensive attempts at taxonomic standardization should be the norm in paleobiological investigations.

The origin of endothermic homeothermy and of high metabolic rate in mammals is currently believed to be the result of early (Mesozoic) selection in advanced cynodont therapsids and/or early mammals for either (1) enhanced thermoregulatory capacity or (2) increased powers of endurance and stamina. Selective factors underlying the origin of specialized respiration/ventilation-support systems in mammals are possible indices of the validity of these two hypotheses. One such support structure is the diaphragm, a specialized muscle that facilitates lung ventilation. We tested capacity for maintenance of resting metabolic rate, thermoregulation, and for extended, intense exercise in laboratory rats (Rattus rattus) in which diaphragm function had been completely ablated. The results were virtual elimination of aeroboic scope (active metabolic rate — resting metabolic rate) but resting metabolic rate was unaffected. Thermoregulatory capacity was unimpaired to at least 8° below lower critical temperature. These and other data suggest that the origin of the mammalian diaphragm, as well as mammalian metabolic rates, may have been related to selection for greater levels of sustainable activity rather than for functions associated with thermoregulation.

Analysis of an exhaustive data base of Namurian ammonoid shell characters indicates that the morphology of the Goniatitida can be explained in terms of functional constraints, resulting particularly from hydrostatic and hydrodynamic properties. Modes of life ranging from benthic to pelagic are inferred on this basis for various goniatitid morphotypes; all morphologic features and facies associations are normally compatible with these inferences. Neutral buoyancy is shown to have been likely for all goniatitids. By contrast, the prolecanitids (Order Agoniatitida) exhibit a number of hydrostatic and morphologic anomalies; these anomalies are not explicable using the same principles and remain problematic. This is noteworthy, in that prolecanitids survived the Permian/Triassic extinctions and gave rise to the diverse ceratitic radiation in the Triassic.

The applicability of these results to ammonoids outside the Namurian is assessed, and, in particular, morphologic parallels with Mesozoic ammonites are discussed.

Like living herbivorous lizards, chelonians, birds, and mammals, plant-eating dinosaurs probably relied on a symbiotic gut microflora, housed in a hindgut fermentation chamber, to break down plant cell wall constituents. Large body sizes in most herbivorous dinosaurs resulted in low mass-specific metabolic rates and low rates of digesta passage through the gut; the effects of large body size were probably enhanced by the low metabolic rates of large dinosaurs as compared with large mammals. The long residence time of digesta in the gut permitted long exposure of refractory plant materials to the microflora, probably enabling even those dinosaurs with unsophisticated dentitions to survive on fodder with high fiber content. Large herbivorous dinosaurs probably fed on plants whose allelochemical defenses were geared more toward reducing digestibility than attacking the herbivore's metabolism directly, obviating the need for a foregut fermentation chamber and permitting these large herbivores to take advantage of the energetic benefits of hindgut fermentation for digestion of low-quality fodder. Differences in dentitions among the groups of herbivorous dinosaurs may correlate with differences in standard metabolic rate, activity level, body size, or food quality, or combinations of these factors, but the relative importance of each is difficult to assess. Because the mass of the fermentation contents was probably large in big herbivorous dinosaurs, the heat of fermentation may have been a significant source of thermoregulatory heat for these reptiles.

The multibrachiate grade of organization in camerate crinoids was principally attained through pinnule differentiation: the accelerated development of a proximal pinnule that ultimately became a pinnule-bearing arm. Although pinnule differentiation is not the only means of achieving a multibrachiate organization in modern crinoids, it was the primary mode in producing both dichotomous and exotomous branching patterns in multibrachiate camerates. Whereas most of the proximal branchings, commonly fixed in the cup of camerates, were due to pinnule differentiaton, the first dichotomy and distal dichotomies above the cup apparently developed from direct modification of a brachial plate at the growing tip of the immature crinoid arm.

Analysis of relative changes in plate position during ontogeny was made on a number of Cenozoic spatangoid echinoids: species of Breynia, Lovenia, Protenaster and Echinocardium. Contrary to the generally held view that adjacent ambulacral and interambulacral columns in echinoids always remain in a fixed relative position during ontogeny, many spatangoids show great fluidity in plate growth, with adjacent columns ‘sliding past’ one another during ontogeny. Furthermore, in many genera plates from one column may undergo strong lateral growth and bisect pairs of plates in adjacent rows. This phenomenon of relative plate movement, both meridional and equatorial, is herein termed ‘plate translocation’. It is considered to occur by localized resorption and redeposition in a narrow zone along adjacent plate boundaries. Plate translocation has been of considerable phylogenetic significance to spatangoids and, combined with an increase in differential allometries between plates, has been one of the most important factors in the evolution of the Spatangoida. Furthermore, the initiation of plate translocation in the apical system in certain Jurassic echinoids may have been the trigger for the migration of the periproct out of the apical system, and a major factor in the evolution of irregular echinoids. Heterochrony in plate growth has been critical in controlling the course of evolution in many lineages of spatangoid echinoids.

Using Sepkoski's compendium of fossil marine families (1982a, and updates), we have analyzed the changing pace of familial origination and extinction within 55 extinct and 44 extant higher taxa of marine organisms. Eight different metrics were calculated, and least-squares regression analysis was used to identify within-taxon trends in the data. All metrics and analyses gave essentially the same results. Origination metrics decline significantly with time during the histories of higher taxa, while extinction metrics increase significantly. The number of statistically significant declines of origination metric, however, substantially and invariably exceeds the number of statistically significant increases of extinction metric for each pair of corresponding metrics analyzed. It follows, therefore, that temporal trends in the pace of origination and extinction within higher taxa are highly asymmetrical.

Further analysis shows that truncating data from temporal endpoints has little effect upon the intensity of origination trends, implying that declining pace of origination is a sustained property of the long term histories of taxa. Such truncation, however, reduces the intensity of extinction trends to statistical insignificance and confirms Van Valen's (1985a) suggestion that extinction behaves largely as a stationary process. If the histories of higher taxa are characterized by substantial declines in the pace of origination while the pace of extinction remains largely stationary, it follows that declining pace of origination is an important controlling factor in long term taxic evolution.