ALBERT

Description: Study of the assemblage of encrusting organisms on co-occurring disarticulated valves of the bivalves Crassostrea virginica and Mercenaria mercenaria in Bogue Sound, North Carolina, indicates that there is little or no substrate specificity among the encrusting organisms but that the shape of the shells has an important influence on how extensively members of each higher taxon collectively inhabit the shells. Encrusting bryozoans, a dense low mat composed of many species from diverse phyla, and a unicellular film cover most of the area of both exterior and interior surfaces. The encrusting bryozoans most extensively cover both surfaces of C. virginica but are in second place behind the multispecies mat on exterior surfaces of M. mercenaria and behind the unicellular film on its interior surfaces. These differences are inferred to result from different physical stability of valves of the two bivalve species, which exhibit different frequencies of circumrotatory growth.Degradation of the assemblage by sodium hypochlorite, to simulate loss of organic matter during fossilization, results in the complete loss of encrusting sponges, erect hydrozoans, erect bryozoans, and ascidians. Loss of these taxa results in overexposure and more apparently uniform distribution of skeletal taxa with respect to their surface representation in living assemblages and also in complete loss of the higher tiers present in the living assemblage. However, indications of the original structural organization of the living assemblage is indicated by preservation of the most abundant taxa in the lower tiers and by the retention in the reduced treated assemblage of the patterns of distribution that characterized the living assemblage.

Description: Bases of attachment of Archimedes colonies are very rare, but many colonies show evidence of origination from a pre-existing set of branches. The evidence consists of spiralled supports (axial screws) originating from fragments of pre-existing sets of branches, proximal taper of colonies or axial screws to a diameter smaller than that of bases of attachment, early stages of new colonies originating from the whorl margins of older colonies, and sets of laterally fused colonies. In the most favorable environments Archimedes populations were dominated by a few genets that consisted of hundreds to perhaps thousands of genetically identical colonies.The ability to reproduce by colony fragmentation was probably an important cause of Archimedes' abundance in the Chesterian seas of eastern North America, especially where it occurred in dense populations in the lee of submarine carbonate shoals. Colony fragmentation, the importance of which is becoming increasingly recognized in modern corals and bryozoans, was also of importance in Paleozoic seas.

Description: Colonies of Late Paleozoic lyre-shaped fenestrate bryozoans began growth on rigid substrates which were on the sea floor. After production of a basal disc, a fenestrate network developed in the shape of an erect bowed fan. The fenestrate fan was initially concave on the apertural side, but the curvature reversed a few millimeters above the base. Colonies were then separated along or just above the basal disc, and rested directly on the sea floor. The zooecial apertures were directed upward. The lateral margin of the fan grew to form the supporting struts of the soon-to-be lyre-shaped colony. An open vault formed below the upwardly bowed fan.Lyre-shaped colonies are interpreted to have lain on the sea floor so that each had the proximal end on the upcurrent side. Feeding zooids were on the upper surface and pulled water through their expanded tentacles. The filtered water passed through the fenestrules, into the vault, and out the open end of the vault, which was on the downcurrent side.

Description: Exploration of the theoretical morphospace of erect helical colony form in Bryozoa, created by McKinney and Raup (1982), reveals that only a small volume of the three-dimensional space of hypothetical form is occupied by actual colonies of the Paleozoic fenestrates (Class Stenolaemata) Archimedes and Helicopora, helical species of the cheilostome (Class Gymnolaemata) Bugula, and the cyclostome (Class Stenolaemata) Crisidmonea archimediformis. Actual helical-colony bryozoans are not found in regions of the morphospace characterized by colony geometries that possess the largest surface areas of filtration sheet. Examination of computer-simulated colonies in the theoretical morphospace reveals that, although possessing high surface areas, colonies in the empty region of high-surface-area morphospace possess other aspects of geometry that are unrealistic as filter-feeding geometries: the filtration-sheet whorls are held at small acute angles to the central colony axis and are deeply nested within one another, both of which are disadvantageous conditions for the system of filter feeding used by the extant cheilostome Bugula, and presumably by extinct helical-colony bryozoans as well.Even though actual bryozoans are found only in the low to intermediate surface-area regions of the theoretical morphospace, surface area of filtration sheet is a major determinant of form in these helical colonies, as is evidenced by a negative correlation in values of the parameters BWANG and ELEV exhibited by the colony data. Minimum values of BWANG are even further constrained by the apparent need of the Archimedes colonies to maintain filtration-sheet branching densities within the range of 20 to 50.

Description: Encrusting bryozoans provide one of the few systems in the fossil record in which ecological competition can be observed directly at local scales. The macroevolutionary history of diversity of cyclostome and cheilostome bryozoans is consistent with a coupled-logistic model of clade displacement predicated on species within clades interacting competitively. The model matches observed diversity history if the model is perturbed by a mass extinction with a position and magnitude analogous to the Cretaceous / Tertiary boundary event. Although it is difficult to measure all parameters in the model from fossil data, critical factors are intrinsic rates of extinction, which can be measured. Cyclostomes maintained a rather low rate of extinction the model solutions predict that they would lose diversity only slowly as competitively superior species of cheilostomes diversified into their environment. Thus, the microecological record of preserved competitive interactions between cyclostome and cheilostome bryozoans and the macroevolutionary record of global diversity are consistent in regard to competition as a significant influence on diversity histories of post-Paleozoic bryozoans.

Description: 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.

Description: Colonies of the fossil bryozoanArchimedesand erect, spiralled species of the living bryozoanBugulaconsist of wedge-shaped systems of radially diverging, bifurcated branches that extend from a helical axial margin. Morphology of these colonies may be simulated using few growth rules. These include 1) radius of the central helical margin (RAD), 2) rate of climb of central helical margin (ELEV), 3) radial angle between successive branches that originate from the central helical margin (ANG), 4) radial growth of all branches, 5) angle between branches and axis of central helical margin (BWANG), 6) distance between three adjacent, radially diverging branches at which the central branch bifurcates into two branches equally spaced between the two side branches (XMIN), and 7) placement of a spacing bar at base of newly bifurcated branches. In addition, size constraints on the simulations must be stipulated.Simulations are begun at a proximal point along the central helix where a radial branch originates and are “grown” in repetitive steps by extending the central helical margin a distance distally, determined by ANG, then censusing established branches for XMIN in order to bifurcate appropriate branches and extend others in several short growth increments, etc. Growth of branches ceases at stipulated maximum width, and growth of the entire simulation ceases at stipulated maximum height.The presence of a helical inner margin marked by uniformly spaced bifurcations generates the spiralled shape, i.e. ELEV must be a positive number. Values of RAD, ELEV (not zero), ANG and BWANG determine form of the spiral; the other growth rules apply to bifurcated unilaminate branch systems in general.The range of observed colony forms and hypothetical morphospace ofArchimedesmay be simulated by varying BWANG, XMIN, and ELEV. RAD was kept constant, as its variation would be redundant with XMIN and ELEV. Variation in ANG affects near-helix morphology, but its influence is undetectable beyond this zone. Variability within colonies may be simulated by assigning to each variable a standard deviation with mean-centered randomly chosen values for each decision.

Description: 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).

Description: Among the various forms developed by marine colonial filter feeders, erect colonial fans have evolved in representatives of several phyla. Branches are relatively narrow, evenly spaced from one another, proliferate by bifurcation, and in most colonies continue from their point of origin to the distal margin of the colony. Their narrowness and uniform placement is apparently related to filling an area with multiple tentacle bells of similar size. The major architectural problem for such a colonial suspension feeder seems to be construction of a sheet composed of narrow branches such that water may be filtered as it flows through the sheet, between branches.The simplest form of the filtration sheets is a uniformly expanding wedge, in which new branches originate at regular intervals from the point of origin of the wedge. Variation in rate of addition of new branches results in more complex forms, such as paraboloids, flaring fans, spirals, and highly folded sheets.

Description: Fusion occurred between pairs of conspecific colonies of Permian fenestrate bryozoans that were of approximately equal radius at time of contact. In contrast, no fusion occurred in pairs of conspecific colonies that were of unequal radius at time of contact. It is suggested that histologically compatible colonies developed from larvae that settled essentially simultaneously and that the larvae were siblings, lecithotrophic with low mobility, and of the same genotype (i.e., polyembryonic). Thus polyembryony appears to have existed as a reproductive “strategy” for at least 275 Myr, and its scarcity in modern taxa cannot be due to recentness of evolution.