Specimens of Peronopora Nicholson, 1881, are abundant in Upper Ordovician rocks of the North American Midcontinent. Based on the positions of units in the Composite Conodont Standard Section, we have sampled 211 specimens over a stratigraphic interval of 9.1 million years. The average duration of sample spacing is 61,664 years but is commonly as small as 32,800 yr.Thirty-four morphometric characters were measured in each specimen and were converted into multistate characters; character-state breaks were established based upon each character's ability to discriminate between phenetic groupings. Each character was subsequently weighted based on the number of derived states, degree of independence from other attributes, and estimated heritability.Cladistic analysis of these data indicate that there are eight species in Peronopora each consisting of an optimally defined crown group and a basal stem group (or paraclade). Character states shared by stem and crown groups define species but, within species, stem and crown groups also differ in some character states. The species are, in ascending order from the base of the tree, Peronopora decipiens (Rominger, 1886), P. compressa (Ulrich, 1979), P. pauca Utgaard and Perry, 1964, P. milleri Nickles, 1905, P. horowitzi new species, P. vera Ulrich, 1888, P. sparsa Brown and Daly, 1985, and finally P. dubia (Cumings and Galloway, 1913). Diagnostic keys permit the unique assignment of each specimen to a species and the separation of members of stem groups from those of crown groups. Thirty-one characters are required to discriminate between all 211 specimens. This contrasts with previous studies of Peronopora where eight or fewer characteristics were used. Of the ten characters most useful in discrimination, only three had been used in the conventional species literature. This accounts, largely, for only 29.8 percent (51 of 171) of previously identified specimens being classified as members of the same species in this analysis. Discriminant function analysis of original measurements, using species identity as the grouping criterion, produces statistically significant separation of species.It appears that stratigraphic position had an explicit and undue effect on previous concepts of species many of which could not be recognized independently of stratigraphic position. All species of Peronopora appear, or are inferred to have appeared, within the Lexington Limestone between the base of the Grier Member and the top of the Millersburg Member. The cladogram indicates that species evolved in a sequential order, but their first appearance datums have been stratigraphically punctuated. Three species have ranges terminating in the Early to Middle Maysvillian, one in the Middle Richmondian, and four in the Late Richmondian. The latter four (or five) of these species died out in the extinction associated with the unconformity at the top of the Richmondian.

The Late Ordovician bryozoan genera of central and southeastern North America are geographically distributed in three biotic provinces, separated by boundaries reflecting major lithofacies differences. The central Cincinnati Province contains most of the North American endemic genera, and represents a narrow ecological zone separating the clastic wedges of the marginal Reedsville‐Lorraine Province from the cratonic carbonate platform of the Red River‐Stony Mountain Province. The provinces provided major life zones, or biomes, for each of the five bryozoan orders. Genera comprising the provinces differed as well in morphologic complexity, geochronologic survivorship, tiering, endemism and eurytopy. Regions on either side of the Cincinnati Province were dominated by inferred immigrants from Baltoscandia. Al‐logenic provincial succession produced time‐averaged mixed faunas in regions near the provincial boundaries. Although most generic originations took place within the Cincinnati Province, evolutionary novelties are associated with the Reedsville‐Lorraine Province. The loss of the diverse Cincinnati Province, connected with global cooling and a eustatic lowering of sea level, may have been a chief factor in the Late Ordovician extinction of bryozoan genera. Genera from the Red River‐Stony Mountain Province differentially survived into the Silurian.

The shape of bryozoan taxonomic survivorship curves is strongly influenced both by grade of morphologic complexity and by mass extinction. Paleozoic bryozoan genera that are morphologically simple have linear taxonomic survivorship; morphologically intermediate taxa have slightly concave survivorship, and complex forms have very concave survivorship. Increasing morphologic complexity, and by inference, increasing specialization of adaptation appear to accompany a systematic departure from a stochastically constant extinction rate. However, the extinctions of the complex taxa are entirely concentrated during three mass extinction events, whereas the extinctions of the simple taxa are more uniformly distributed throughout the Paleozoic; the extinction pattern of the morphologically intermediate taxa is intermediate to those of the simple and complex groups. Exclusion of the genera affected by mass extinction increases the convexity of the survivorship curves, and reverses the apparent correlation of extinction rate with morphologic complexity. The macroevolutionary pattern of the complex genera resembles an r-strategy, whereas that of the simple taxa resembles a K-strategy.

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.

A paradoxical relationship exists between the genetic and morphologic adaptive strategies of benthic marine invertebrates; morphologically variable species from unstable environments have been shown to possess less genetic variability than species with more constant phenotypes from stable habitats. The mode of growth of Ordovician bryozoans provides an insight into this paradox. These bryozoans exhibit morphologic gradients within zooidal subcolonies that change throughout colony development. Entire clusters of zooids begin and cease growth as a function of their spatial position with respect to neighboring clusters. A comparison of the within-colony and among-colony components of developmental and morphologic variability was made from populations inhabiting Ordovician environments of differing stability. In four stratigraphically persistent species, the level of developmental variability is homogeneous within species, but varies significantly across taxa. The two species with the highest levels of developmental variability (less canalized development) fit the concept of r-selected opportunistic species and are most abundant in communities of lowest diversity. The other two species have much lower levels of variability (more canalized development), fit the concept of K-selected equilibrium species, and are most abundant in the communities of highest diversity. Within-colony morphologic variability is also higher in the opportunistic rather than the equilibrium species, indicating that the higher morphologic variability observed in unstable environments is the product of within-genotype deregulation, and not the result of higher genetic polymorphism. The equilibrium species in stable environments have lower levels of morphologic deregulation and correspondingly greater variation among genotypes than within genotypes in fossil populations.