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.

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.

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.

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.

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).

One of the striking yet scarcely documented episodes of clade replacement in the post-Paleozoic fossil record is the decline of cyclostome Bryozoa and the corresponding, rapid diversification of cheilostome Bryozoa. These clades are closely associated morphologically and phylogenetically, and their ecological similarities have previously led to the inference that competition was a primary cause of the overt pattern of replacement. Alternatively, previous compilations of bryozoan families and genera have implied that extinctions at the Cretaceous/Tertiary boundary differentially affected cyclostomes, and thus were also an important factor in the transition.

We first evaluated the ecological context for competition between the two clades, then updated and reexamined the history of absolute family diversity for bryozoans in consecutive geologic stages from Jurassic to Recent. The resulting trends echo the patterns shown in earlier family level compilations, but indicate a slight shift in the frequency of cheilostome family originations from Late Cretaceous to early Paleogene. The relative fall in cyclostome family diversity at the Cretaceous/Tertiary boundary is significantly less than shown in earlier genus level compilations. We then assessed these various compilations of absolute diversity by analyzing species counts and percentages in 728 fossil assemblages, primarily from North America and Europe, over the same time interval. Cyclostome species overwhelmingly dominate assemblages from Jurassic through Cenomanian, then decline significantly in average percentage dominance through the Campanian. Cheilostomes are predominant in Campanian and later assemblages. Cyclostome species percentages do decrease overall through the Tertiary, but this decrease is small and non-uniform, varying around 30%, with a sharp drop in the Late Neogene. Our within-assemblage results indicate that as cheilostomes radiate, their mean species diversity, maximum diversity, and variance all increase, thereby accounting for much of the decline in average percentage of cyclostomes within assemblages. While this result does not exclude a role for competition, an hypothesis of relative decline in cyclostome species richness based on competitive extinction alone seems unlikely. Further, despite decreases in absolute species counts following end-Cretaceous extinctions, within-assemblage percentages of cheilostome or cyclostome species show only slight change relative to one another. Comparison of these and earlier diversity compilations indicates that the dynamics of bryozoan clade replacement may be perceived differently at different ecologic scales or taxonomic ranks.

The biomass of marine consumers increased during the Phanerozoic. This is indicated by the increase in both fleshiness and average size of individuals of dominant organisms, coupled with the conservative estimate that dominant organisms in the Cenozoic are at least as abundant as those in the Paleozoic. As faunal dominants replaced one another during the Phanerozoic the general level of metabolic activity increased due to both increase in basal metabolism and increase in more energetic modes of life. This demonstrates that the expenditure of energy by marine consumers has increased with time as well. There is a time lag in the expansion of more energetic life habits from environmental settings known to have high food supply into regions expected to have lower rates of food supply (e.g., bivalves into offshore carbonate environments or deep burrowing deposit feeders into the full range of shelf environments), and a time lag in diversification of energetic modes of life (e.g., predation or deep burrowing deposit feeding) for long intervals after they first appeared. This suggests that the supply of food increased across the whole spectrum of marine habitats during the Phanerozoic. The great diversification of specialized predators especially suggests that biomass increase took place all the way down the food chain to the level of primary production. The development of plant life on land and the impact of land vegetation on stimulating productivity in coastal marine settings, coupled with the transfer of organic material and nutrients from coastal regions to the open ocean, and the increase through time in diversity and abundance of oceanic phytoplankton all point to increased productivity in the oceans through the Phanerozoic.