A simple equilibrial model for the growth and maintenance of Phanerozoic global marine taxonomic diversity can be constructed from considerations of the behavior of origination and extinction rates with respect to diversity. An initial postulate that total rate of diversification is proportional to number of taxa extant leads to an exponential model for early phases of diversification. This model appears to describe adequately the “explosive” diversification of known metazoan orders across the Precambrian-Cambrian Boundary, suggesting that no special event, other than the initial appearance of Metazoa, is necessary to explain this phenomenon. As numbers of taxa increase, the rate of diversification should become “diversity dependent.” Ecological factors should cause the per taxon rate of origination to decline and the per taxon rate of extinction to increase. If these relationships are modeled as simple linear functions, a logistic description of the behavior of taxonomic diversity through time results. This model appears remarkably consistent with the known pattern of Phanerozoic marine ordinal diversity as a whole. Analysis of observed rates of ordinal origination also indicates these are to a large extent diversity dependent; however, diversity dependence is not immediately evident in rates of ordinal extinction. Possible explanations for this pattern are derived from considerations of the size of higher taxa and from simulations of their diversification. These suggest that both the standing diversity and the pattern of origination of orders may adequately reflect the behavior of species diversity through time; however, correspondence between rates of ordinal and species extinction may deteriorate with progressive loss of information resulting from incomplete sampling of the fossil record.

A kinetic model of Phanerozoic taxonomic diversity I. Analysis of marine orders

Malformations of trilobites are classified as healed injuries, teratological conditions, and pathological conditions. An improved method of recognizing such malformations combines information about the conditions under which cell injury can occur, the processes by which animal tissues react to injury, and trilobite morphology. Study of healed injuries of polymeroid trilobites shows that injuries attributed to sublethal predation tend to be most commonly preserved on the pleural lobes, the posterior half of the body, and the right side. Statistically significant differences in the number of predation scars between the right and left sides is interpreted as evidence of right-left behavioral asymmetry in some predators of trilobites or the trilobites themselves. Asymmetrical, or lateralized, behavior in present-day animals is one manifestation of handedness, and is usually related to a functional lateralization of the nervous system. Evidence of behavioral lateralization in some Paleozoic predators or prey suggests that those organisms also possessed lateralized nervous systems. Right-left differences in preserved predation scars on trilobites date from the Early Cambrian (Olenellus Zone), and are the oldest known evidence of behavioral asymmetry in the fossil record. Other examples of structural or behavioral asymmetry from the fossil record of animals are cited. Lateralization is recognized in representatives of the Arthropoda, Annelida, Bryozoa, Echinodermata, Cnidaria, Mollusca, Chordata, and Conodonta, and in trace fossils.

Trilobite malformations and the fossil record of behavioral asymmetry

Limestone beds in the Middle Ordovician (Trentonian) Bobcaygeon Formation, exposed near Kirkfield, Ontario, exhibit irregular, bored and encrusted surfaces indicative of early lithification. These hardgrounds were formed in extensively burrowed carbonate sediments and their hummocky surface topographies were inherited, in part, from a pre-existing pattern of burrow tunnels. A diverse community, including bryozoans, brachiopods, crinoids, edrioasteroids, and paracrinoids, colonized these hard substrates. In addition, most surfaces are riddled with small, cylindrical boreholes (Trypanites) which represent dwellings of soft-bodied organisms. Some hardground surfaces were inhabited by multiple generations of organisms. Remains of the older generations of encrusters were strongly abraded and nearly obliterated. Superimposed upon these worn remnants are well-preserved remains of rather fragile organisms, e.g. complete hybocystitid crinoids and edrioasteroids. Evidently, certain hardground surfaces were rapidly buried by muds, resulting in the in situ preservation of the last generation of attached organisms.Slightly differing subcommunities of organisms inhabited various microhabitats provided by the irregular hardground surfaces. Thus, the relative abundance of bryozoans and echinoderms encrusting the roofs of small crevices differs from that on the exposed upper surfaces of the hardgrounds. This is the geologically oldest known hardground community in which microhabitat subdivision can be recognized. However, polarity between the subcommunities is not as pronounced as in geologically younger hard substrate faunas.

Preservation and paleoecology of a Middle Ordovician hardground community

Echinoderms have long been characterized by the presence of ambulacra that exhibit pentaradiate symmetry and define five primary body axes. In reality, truly pentaradial ambulacral symmetry is a condition derived only once in the evolutionary history of echinoderms and is restricted to eleutherozoans, the clade that contains most living echinoderm species. In contrast, early echinoderms have a bilaterally symmetrical 2–1–2 arrangement, with three ambulacra radiating from the mouth. Branching of the two side ambulacra during ontogeny produces the five adult rays. During the Cambrian Explosion and Ordovician Radiation, some 30 clades of echinoderms evolved, many of which have aberrant ambulacral systems with one to four rays. Unfortunately, no underlying model has emerged that explains ambulacral homologies among disparate forms. Here we show that most Paleozoic echinoderms are characterized by uniquely identifiable ambulacra that develop in three distinct postlarval stages. Nearly all “aberrant” echinoderm morphologies can be explained by the paedomorphic ambulacra reduction (PAR) model through the loss of some combination of these growth stages during ontogeny. Superficially similar patterns of ambulacral reduction in distantly related clades have resulted from the parallel loss of homologous ambulacra during ontogeny. Pseudo-fivefold symmetry seen in Blastoidea and the true fivefold symmetry seen in Eleutherozoa result from great reduction and total loss, respectively, of the 2– 1–2 symmetry early in ontogeny. These ambulacral variations suggest that both developmental and ecological constraints affect the evolution of novel echinoderm body plans.

Ontogeny in the fossil record: diversification of body plans and the evolution of “aberrant” symmetry in Paleozoic echinoderms

A new edrioasteroid fauna from the Ordovician of Morocco, North Africa includes eight new isorophid species placed in six genera of which three are new. A phylogenetic analysis of Moroccan genera and other taxa places these new edrioasteroids into a phylogenetic framework. Pyrgocystidae is redefined to include plesiomorphic isorophids with hood plates underlying the cover plates. New pyrgocystid taxa include Streptaster nodosus sp. nov., Belochthus? chauveli sp. nov., Argodiscus espilezorum sp. nov. and Moroccopyrgus matacarros gen. et sp. nov. Isorophinid taxa include Isorophus africanus sp. nov., Isorophusella gutii sp. nov., Euryeschatia reboulorum gen. et sp. nov. and the morphological aberrant Anedriophus moroccoensis gen. et sp. nov. These taxa are attached either epibiotically on conulariids or echinosphaeritid rhombiferans, or encrust large bioclasts such as cephalopods and trilobites in a siliciclastic environment that is otherwise devoid of suitable hard substrates. The presence of close relativ...

Ordovician edrioasteroids from Morocco: faunal exchanges across the Rheic Ocean

A Study Of North American Edrioasteroidea.

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A Study Of North American Edrioasteroidea.