Evolution does not occur in a vacuum. It takes place against a background of changing conditions, some of a climatic or tectonic nature, some created by organisms themselves. The extent to which the rate and direction of evolution are controlled by organisms is the central question in a debate that has been raging for as long as evolution has been part of the intellectual vocabulary of scientists. In an effort to stimulate empirical work on the subject, we organized a symposium on the contributions that functional morphology and comparative physiology are making to paleobiology. The symposium was held as part of the North American Paleontological Convention on July 1, 1992, at the Field Museum of Natural History in Chicago. This issue of Paleobiology contains all the submitted and accepted papers presented at the symposium.

Approaches to patterns of diversification based on counting taxa at a given rank can be misleading, even when all taxa are monophyletic. Such “rank-based” approaches are unable to reflect a hierarchy of evolutionary events because taxa of the same rank cannot be nested within one another. Phylogenetic trees specify an order of origination of characters and clades and can therefore be used in some cases to test hypotheses on causal relationships between characters and changes in diversity. “Tree-thinking” also clarifies discussions of the age of groups, by distinguishing between splitting of the stem-lineage from its sister group and splitting of the crown-group into extant clades.

Cladistic evidence that Pentoxylon, Bennettitales, and Gnetales are the sister group of angiosperms implies that the angiosperm line (angiophytes) existed by the Late Triassic. The presence of primitive members of five basic angiosperm clades indicates that the crown-group (angiosperms) had begun to diversify by the mid-Early Cretaceous (Barremian-Aptian), but not necessarily much earlier. The greatest unresolved issue raised by cladistic analyses concerns the fact that the angiosperm tree can be rooted in two almost equally parsimonious positions. Trees rooted near Magnoliales (among “woody magnoliids”) suggest that the angiosperm radiation may have been triggered by the origin of intrinsic traits, e.g., a fast-growing, rhizomatous habit in the paleoherb and eudicot subgroup. However, trees rooted among paleoherbs, which are favored by rRNA data, imply that these traits are basic for angiosperms as a whole. This could mean that the crown-group originated not long before its radiation, or, if it did originate earlier, that its radiation was delayed due to extrinsic factors. Such factors could be a trend from environmental homogeneity and stability in the Jurassic to renewed tectonic activity and disturbance in the Early Cretaceous. Potentially relevant pre-Cretaceous fossils cannot be placed with confidence, but may be located along the stem-lineage (stem angiophytes); their generally paleoherb-like features favor the paleoherb rooting. The history of angiophytes may parallel that of Gnetales: some diversification of the stem-lineage in the Late Triassic, near disappearance in the Jurassic, and vigorous radiation of the crown-group in the Early Cretaceous.

During the latest Cretaceous and the Paleocene in western North America, disappearance rates for mammalian genera track appearance rates, both reaching their peak in the early Paleocene (Puercan) following the extinction of non-avian dinosaurs. Some of the disappearances during this time were pseudoextinctions that resulted when ancestral species disappeared during speciation.

Species-level cladistic analyses and a well-constrained biostratigraphic framework are required to study this form of pseudoextinction. Cladistic analyses show that monophyly cannot be established or rejected for some species because these species lack autapomorphies (uniquely derived character states) that unite their constituent members. Such taxa, termed metaspecies, are potential ancestors to species and higher clades with which they share a node in the cladogram.

A hypothetical species-level cladistic analysis coupled with three different hypothetical biostratigraphies shows how different models of speciation (bifurcation, budding, or anagenesis) result in very different patterns of true versus pseudoextinction. Depending on the speciation model, true extinction can be overestimated by as much as a factor of four, raising the specter of mass extinction. Species-level studies for three early Tertiary mammalian taxa—taeniodont eutherians, taeniolabidid multituberculates, and periptychid ungulates—use the same procedures. They show that almost 25% of disappearances during the early Paleocene (Puercan) for species in the analysis were pseudoextinctions of metaspecies. Budding and anagenetic-like peripatric speciation, but not bifurcation, are seen in the three examples.

Equating disappearance to true extinction can profoundly affect interpretations of faunal turnover, especially during mass extinctions or major faunal reorganizations. Some authors use pseudoextinction to describe the taxonomic rather than evolutionary disappearance of nonmonophyletic groups. Pseudoextinction, as used here refers only to the evolutionary disappearance of metaspecies via speciation. Both usages seem appropriate but should not be confounded.

The problem of how accurately paraphyletic taxa versus monophyletic (i.e., holophyletic) groups (clades) capture underlying species patterns of diversity and extinction is explored with Monte Carlo simulations. Phylogenies are modeled as stochastic trees. Paraphyletic taxa are defined in an arbitrary manner by randomly choosing progenitors and clustering all descendants not belonging to other taxa. These taxa are then examined to determine which are clades, and the remaining paraphyletic groups are dissected to discover monophyletic subgroups. Comparisons of diversity patterns and extinction rates between modeled taxa and lineages indicate that paraphyletic groups can adequately capture lineage information under a variety of conditions of diversification and mass extinction. This suggests that these groups constitute more than mere “taxonomic noise” in this context. But, strictly monophyletic groups perform somewhat better, especially with regard to mass extinctions. However, when low levels of paleontologic sampling are simulated, the veracity of clades deteriorates, especially with respect to diversity, and modeled paraphyletic taxa often capture more information about underlying lineages. Thus, for studies of diversity and taxic evolution in the fossil record, traditional paleontologic genera and families need not be rejected in favor of cladistically-defined taxa.

Quantitative data on lycopsid and aquatic fern megaspore taxa recovered from Carboniferous, Mesozoic, and Tertiary strata have been compiled in order to analyze the changes in diversity of the two groups of fossil plants that produced them. Numbers of species of lycopsid megaspores are similar in the Carboniferous and Mesozoic, whereas the diversity of megafossils is much lower in post-Paleozoic deposits. Our data suggest that lycopsids were more diverse in the Mesozoic than previously thought and that there is a preservational bias against the megafossils, because the plants were probably mainly herbaceous. Heterosporous aquatic ferns first appeared in the Neocomian and gradually diversified until the early Late Cretaceous, after which their numbers remained relatively stable, whereas the variety of lycopsids declined dramatically during the Late Cretaceous. These changes occurred at a time of rapid angiosperm diversification. The reduced diversity of the lycopsids may have been caused by the invasion of their aquatic and damp forest-floor habitats by heterosporous ferns and by aquatic and herbaceous angiosperms. These diversity changes do not seem to be directly related to the global events at the Cretaceous-Tertiary boundary, but the relatively few samples available and the resulting range truncation would make detection of such correlations difficult.

The compound eyes of trilobites provide the best examples of fossilized sensory organs for which the function in life can be worked out today because the optical array of their corneal lenses preserves the geometry with which the eye originally sampled the visual world. An analysis of trilobite vision is strengthened by the use of new mathematical approaches to compound eye design. In particular, the product of the facet diameter (D) and the interommatidial angle (Δϕ) gives the value of the eye parameter, DΔϕ, which is a reliable indicator of the photic conditions in which the eye was used. In modern arthropods, DΔϕ values range from 0.3 for animals active in bright sunlight to 20 or more for nocturnal or deep-sea animals. Two major types of compound eyes existed in trilobites: schizochroal and holochroal. In our previous work with schizochroal eyes in the phacopids Phacops rana crassituberculata and Phacops rana milleri, we found that eye parameter values ranged from 10 to >150. These values of the eye parameter are much greater than in any living arthropod, implying that modern compound eye theory does not apply to schizochroal eyes. We suggested that each ommatidium of the schizochroal eye served as a miniature lens eye. If so, phacopid vision must have been unique, with multiply overlapping visual fields. In the new work of this paper, we examined holochroal compound eyes in Asaphus cornutus, Isotelus gigas, and Homotelus sp. Holochroal eyes contain far more ommatidia than do schizochroal types, reducing both facet diameter (D) and interommatidial angle (Δϕ). Thus, DΔϕ values in these species fall into the same range as in modern nocturnal compound eyes. This implies that function of the holochroal eye was similar to that of modern arthropods, and that they were used in moderate to dim intensities of light.

Two methods are discussed for assessing the contributions of subgroups to the morphological disparity of the larger group containing them. (1) Given an ordination of points representing specimens or species in morphological space, morphological disparity of the entire group is measured as the average squared distance of points from the centroid. The contribution that a subgroup makes to morphological disparity is measured as the average squared distance of its points from the overall centroid (not the subgroup centroid), weighted by the subgroup sample size relative to the total group sample size. Thus, morphological disparity of a group can be additively partitioned into the disparity components of its subgroups, and the relative contributions of these subgroups can be assessed quantitatively. (2) An alternative approach is to compare morphological disparity of a group to the disparity it would have if a certain subgroup were omitted. If the resulting disparity differs substantially from the original disparity, then the subgroup in question is considered to have a significant effect on morphological disparity. Because some subgroups are very centralized in morphological space, omitting them can cause an increase in morphological disparity when disparity is measured as the average dissimilarity among species. In general, relatively large subgroups that are located peripherally in morphospace make the greatest contributions to morphological disparity, and failure to sample smaller groups often has little effect on disparity estimates. The two methods are applied to morphological disparity in trilobites, partitioned at different levels in the taxonomic hierarchy. Results of the two methods are intuitively reasonable and largely in agreement, and point to the predominance of Early Cambrian olenelloids, Cambro-Ordovician Libristoma, Ordovician Asaphina and Cheirurina, Siluro-Devonian Phacopida and Phacopina, and Devonian Proetida.

Rapid shell closure in articulate brachiopods occurs by a twitch contraction of the “quick” adductor muscles. Rapid accelerations and decelerations of the valves induce large accelerations of the surrounding fluid, placing constraints on the speeds of closure; inertial forces govern the mechanics of the closing mechanism. The position, size, and physiological properties of the muscles, and the size and shape of the shell, govern the properties of the shell closing system. A numerical model predicts that, for given muscle and shell characteristics, there is a location of the adductor muscle which maximizes the shell's closing velocity, and that this position shifts allometrically with size. Positive allometry in the muscle's moment arm length is observed in both living and fossil brachiopods, although the observed coefficients are less than those predicted by the model.

The brachiopod closing mechanism provides a model for understanding the biomechanical interrelationships between components of dynamic skeletal systems. Dynamic systems, where the muscles induce resisting inertial reactions, are characterized by mechanical feedback loops, or couplings. Mechanical couplings may represent an important class of constraints on the evolution of skeleto-muscular architecture.

A comparison is made between compilations of times of origination and extinction of fossil marine animal families published in 1982 and 1992. As a result of ten years of library research, half of the information in the compendia has changed: families have been added and deleted, low-resolution stratigraphic data have been improved, and intervals of origination and extinction have been altered. Despite these changes, apparent macroevolutionary patterns for the entire marine fauna have remained constant. Diversity curves compiled from the two data bases are very similar, with a goodness-of-fit of 99%; the principal difference is that the 1992 curve averages 13% higher than the older curve. Both numbers and percentages of origination and extinction also match well, with fits ranging from 83% to 95%. All major events of radiation and extinction are identical. Therefore, errors in large paleontological data bases and arbitrariness of included taxa are not necessarily impediments to the analysis of pattern in the fossil record, so long as the data are sufficiently numerous.

Morphological and taxonomic diversity each provide insight into the expansion and contraction of major biological groups, while the nature of the relationship between these two aspects of diversity also has important implications for evolutionary mechanisms. In this paper, I compare morphological and taxonomic diversity within the classes Blastoidea and Trilobita, and within the trilobite clades Libristoma, Asaphina, Proetida, Phacopida, and Scutelluina. Blastoid morphology is quantified with homologous landmarks on the theca, and trilobite form is measured with a Fourier description of the cranidium. Morphological diversity is measured as the total variance among forms in morphological space (proportional to the mean squared distance among forms). Blastoid taxonomic diversity is based on published compilation of stratigraphic ranges of genera. The Zoological Record was used to determine the number of new species of trilobites described since the publication of the Treatise; temporal patterns in species richness are similar to those for generic richness based on the Treatise, suggesting a common underlying signal.

Morphological variety and taxonomic richness often increase together during the initial diversification of a clade. This pattern is consistent with diffusion through morphospace, although some form of adaptive radiation cannot be ruled out. Morphological diversity varies little throughout much of the history of Proetida, a pattern that may suggest major constraints on the magnitude and direction of evolution, and that agrees with the perception of Proetida as a morphologically conservative group. Two major patterns are seen during the decline of clades. In Blastoidea, Trilobita, Libristoma, and Asaphina, morphological diversity is maintained at substantial levels, and in fact continues to increase, even in the face of striking reductions in taxonomic richness. This pattern suggests continued diffusion through morphospace and taxonomic attrition that is effectively non-selective with respect to morphology. In Phacopida, Scutelluina, and to some extent in Proetida, morphological diversity decreases along with taxonomic diversity. This pattern suggests heterogeneities such as elevated extinction and/or reduced origination in certain regions of morphospace. As found previously for the echinoderm subphylum Blastozoa, all studied clades of trilobites except Proetida show maximal morphological diversity in the Mid–Late Ordovician and maximal taxonomic diversity sometime during the Ordovician, suggesting some degree of common control on diversification patterns in these groups.

The evolutionary rates of Paleozoic Crinoidea were obtained using dynamic survivorship analysis. The stratigraphic ranges of 838 genera were used in the analyses, revealing a mean generic duration of 12.0 m.y. and a mean species duration of 6.7 m.y., values within the range of longevities reported for other taxa.

Further analyses showed differences in evolutionary rates among crinoid taxa: camerate species and genera were shorter-lived than species and genera of flexibles and inadunates. This pattern may result from ecological differences among these taxa: an energy budget equation solved for crinoids with various filter morphologies revealed that crinoids with fine-mesh filters require higher current velocities to supply them with sufficient particulate nutrients than do crinoids with coarse-mesh filters. A hypothesis stipulating that these differences control the distribution of crinoids among different environments is supported by patterns of occurrence of Mississippian crinoids: the pinnulate camerates (fine filter) dominate higher energy settings while the non-pinnulate inadunates and flexibles (coarse filter) are found in all environments. The “specialized” pinnulate crinoids may therefore be more prone to speciation and extinction than the non-pinnulate “generalists,” thus accounting for the observed differences in the evolutionary rates of the three subclasses.

The above hypothesis was tested by comparing evolutionary rates of two morphological groups: fine-filtered crinoids (camerates) and coarse-filtered crinoids (non-pinnulate Paleozoic crinoids). As predicted, fine-filtered taxa had higher extinction and origination rates. A “bootstrapping” technique revealed that the differences in extinction rates were significant at the p < 0.10 level.

We studied mammal and bird track formation at the northern edge of Lake Manyara, Tanzania, to develop models for interpreting fossil tracks and trackways. Lake Manyara is a closed-basin, alkaline lake in the East African Rift System. The area has a high vertebrate diversity, allowing us to investigate tracks in an environment similar to that of many ancient track-bearing sequences. Three study sites, two on mud flats adjacent to the lake margin and a third on a delta floodplain, provided contrasting environments in which to assess the types of biological data that can potentially be extracted from fossil trackways.

Our censuses of mammals and their tracks revealed that most species that occur within the study area leave a track record, and that common species leave abundant tracks, although numbers of trackways are not proportional to numbers of individuals. Logarithmic increases in track sampling area yield a linear increase in the proportion of both the medium and large-sized local mammals represented in a track record. Transect vs. area mapping methods produced different censusing results, probably because of differences in monitoring periods and areal coverage.

We developed a model of expected track production rates that incorporates activity budget and stride length data in addition to abundance data. By using these additional variables in a study of diurnal birds, we obtained a much better estimator relating track abundance to trackmaker abundance than that provided by census data alone. Proportions of different types of tracks predicted by the model differ significantly from the observed proportions, almost certainly because of microenvironmental differences between the censusing and track counting localities. Censuses of fossil tracks will be biased toward greater numbers of depositional-environment generalists and away from habitat-specific species.

Trackways of migratory animals were dominantly shoreline-parallel, whereas trackways of sedentary species were more variable. A strong shoreline-parallel environmental zonation at the Alkaline Flats site exerted an influence on trackmaker distribution patterns, initial track formation, and track preservation. Variations in habitat usage by different species, as well as species abundance and directionality of movement, were all important in determining the number of preservable tracks a species produced within a given environmental zone.

Fossil trackways are time-averaged, although over entirely different temporal scales than are bones. Unlike bones, tracks are not space-averaged. Therefore, wherever possible, fossil track and bone studies should be used to complement each other, as they provide fundamentally different pictures of paleocommunities. Tracks provide “snapshot” views of localized assemblages of organisms useful in reconstructing autecological relationships, whereas bones yield a broader image of a local fauna in which seasonal and microenvironmental variation are more commonly smoothed out.

This analysis examines the evolution of the greater diversity of species with non-planktonic larval types relative to species with planktonic larval types in the turritellid gastropods. This sort of trend has been documented in both the fossil and recent biota of several gastropod families. Two mechanisms for generating diversity gradients in larval types have been proposed in the literature. The first, species selection, focuses on the population biology of larval types. The second proposes that factors in development that are mediated by organismal adaptation are responsible. Turritellids have been cited as a classic example of species selection. In order to examine the relevance of these two proposed mechanisms, a phylogenetic analysis of the turritellids using molecular sequence data was performed to determine the evolution of larval types in this clade. The resultant phylogeny suggests that species selection is not the only process driving the trend toward increasing numbers of non-planktonic species through time. Developmental processes, apart from those involving organismal adaptation (except in the trivial sense), are implicated as playing a role in this trend. In particular, these processes may involve changes in the timing of germ-line sequestration in organisms. Germ-line sequestration governs how accessible organisms are to heritable variation during ontogeny. Embryological evidence from gastropods suggests that non-planktonic species have early germ-line sequestration relative to planktonic species, making them more resistant to developmental change. Thus, non-planktonic lineages will only rarely revert to a planktonic larval mode.

This paper presents scaling equations relating suspension-feeding rates to body size for articulate brachiopods and bivalve molluscs, two classes which represent a significant component of the fossil record of marine benthic communities. Clearance (feeding) rates of five species of living articulate brachiopods and three species of epifaunal suspension-feeding bivalve molluscs collected from mid-latitude fjords of Newfoundland and New Zealand were measured in similar experimental conditions. In comparisons within and between the two classes, we found that both plectolophous and spirolophous brachiopods had significantly lower feeding rates than mytilids, which are filibranchs, but that a sympatric primitive eulamellibranch veneroid bivalve had rates comparable to the brachiopods. Articulate brachiopods do not appear to feed effectively at the high algal concentrations which bivalves can exploit. The data on comparative suspension-feeding rates support the hypothesis that past changes in diversity and distribution of bivalves and brachiopods may be related to an overall increase in energy flux and escalation of metabolic rates during the Phanerozoic.

Cladistic analysis of osteological and dental characters in a monophyletic group of Miocene and younger tayassuids demonstrates a pattern of changes in the degree of sexual dimorphism in canine tooth diameter and zygomatic arch width, and in phenotypic correlations between these characters. Primitively, tayassuids have canine teeth that are sexually dimorphic and discretely bimodal in size, and zygomatic arches that are narrow in both sexes. Many late Miocene and Pliocene tayassuids have broad, winglike zygomatic processes. In some species, these processes are large in both sexes, but in others, those of females are much smaller than those of males. The presence of large processes in both sexes is primitive relative to the condition of strong sexual dimorphism. In five separate clades, the zygomatic processes of both sexes become reduced in size, and the degree of sexual dimorphism in canine size becomes reduced as well. The pattern is congruent with predictions derived from a theoretical model of the evolution of sexual dimorphism, and it further indicates the emergence of a new phenotypic correlation between two previously uncorrelated characters, canine size and zygoma size. The advent of this new correlation coincides with the advent of pronounced sexual dimorphism in zygomatic processes. Although such a pattern could be explained by genetically modifying phenotypic expression of homologous characters in one sex or the other, an epigenetic modification of expression is equally plausible: the evolution of sexual dimorphism in homologous characters could be accomplished by placing phenotypic expression of an originally monomorphic character under the control of steroid sex hormones. This hypothesis is consistent with evidence from many vertebrate groups, and it provides testable predictions.

In this paper we analyze the laws of growth that control planktic foraminiferal shell morphology. We assume that isometry is the key toward the understanding of their ontogeny. Hence, our null hypothesis is that these organisms construct isometric shells. To test this hypothesis, geometric models of their shells have been generated with a personal computer. It is demonstrated that early chambers in log-spirally coiled structures cannot follow a strict isometric arrangement. In the real world, the centers of juvenile chambers deviate from the logarithmic growth curve. Juvenile stages are generally more planispiral and contain more chambers per whorl than adult stages. These traits are shown to be essential in order to keep volumes of consecutive chambers in geometric progression. We are convinced that the neanic stage marks the constructional bridge from a juvenile set of growth parameters to an adult one. The adult stage can be strictly isometric, that is, the effective shape is constant and the increase in volume after a chamber addition is proportional to the preexisting volume of the shell.

The shell volume is related to the biomass, the ratio of outer shell surface area to shell volume is related to the respiration rate and the ratio of the total shell surface area to shell volume is related to the total calcification effort. The influence of the parameters of the model on these relationships is investigated. Only the initial radius and the rate of radius increase affect the relationships between shell volume and surface area. The other shape parameters merely provide a fine tune-up of these relationships. Size itself plays a major role during foraminiferal development.

The 18 m.y. history of the subfamily Equinae (exclusive of Archaeohippus and “Parahippus”) in North America consisted of a 3-m.y. radiation phase, a 9-m.y. steady-state diversity phase, and a 6-m.y. reduction phase. During the steady-state phase, species richness varied between 14 and 20, with two maxima at about 13.5 and 6.5 Ma. Species richness of the tribes Hipparionini and Equini was about equal through the middle Miocene, but hipparionines consistently had more species in the late Miocene and early Pliocene. Overall mean species duration was 3.2 m.y. (n = 50), or an average extinction rate of 0.31 m.y.-1 During the radiation phase, speciation rates were very high (0.5 to 1.4 m.y.-1), while extinction rates were low (<0.10 m.y.-1). Speciation and extinction rates both averaged about 0.15 m.y.-1 during the steady-state phase, with extinction rates having more variation. Extinction rates increased fourfold during the reduction phase, while speciation rates declined slightly. Late Hemphillian extinctions affected both tribes severely, not just the three-toed hipparionines, and were correlated with global climatic change.

Pterobranchs are the closest living relatives of graptolites. Their skeleton is constructed from the same material, and in a homologous manner. Growth rates of the pterobranch Cephalodiscus gracilis are described for the first time and, along with rhabdopleuran growth rates, they are used to estimate the amount of time invested by a graptolite colony in growing its rhabdosome. This is a measure of minimum age. The significance of age calculations is shown for individuals and large communities of graptoloids. Large individuals can be shown to be much older than the time it would have taken them to settle through seawater and so it is shown that graptoloids moved up, as well as down, through the water column. Life tables constructed for biserial graptoloids from the Utica shale in Quebec, Canada, suggest that these graptoloids died from constant environmental stress. Graptoloid length can thus be a function of environment and should only cautiously be considered to be of taxonomic significance.

Zooids of cheilostome bryozoans are on average substantially more robust than are zooids of cyclostome bryozoans. The differences include greater number, length, and cross-sectional area of tentacles, plus a more extensively developed funiculus. Median values for mouth size and cilia-generated feeding current velocity are greater for cheilostomes than for cyclostomes so that cheilostomes have the potential for greater intake of nutrient energy per unit time, which may explain their apparently higher growth rates. For unit area of substrate occupied, the Cheilostomata (Class Gymnolaemata; members of the post-Paleozoic fauna) contain greater biomass and apparently generate greater energy flow than do the Cyclostomata, which are the only extant order of the Class Stenolaemata (characteristic of the Paleozoic fauna).