6/15/2015 GLY3603 Paleontology Module 4 – Lecture 4 Life of the Ordovician. Changes in Life Through Time: The Paleozoic Era A. Invertebrate Evolutionary Trends 1. Mobility to Stability. • Cambrian assemblages were mostly mobile forms of metazoans, trilobites the most abundant of them. • The Ordovician Radiation included organisms that were sessile and had hard parts. • An Early Ordovician change in seawater chemistry facilitated precipitation of calcite over aragonite (a less stable mineral) from seawater and the formation of widespread “hardgrounds” for invertebrate attachment. A. Invertebrate Evolutionary Trends Continued 2. Increased diversity and the rise of the “Paleozoic Fauna” • • • • • • A great adaptive radiation of invertebrate fauna. Number of families of invertebrates increased from about 150 at the end of the Cambrian to 400 during the Ordovician. The Cambrian fauna was replaced by a fauna that persisted throughout the rest of the Paleozoic, a period of more than 230 million years. Epifaunal suspension feeders, especially bryozoans, brachiopods, corals and stalked echinoderms, dominated this fauna, the “Paleozoic Fauna”. Infauna included suspension feeders in shallow burrows and deposit feeders Epifaunal deposits feeders, scavangers, grazers, parasites and predators also composed the community 1 6/15/2015 A. Invertebrate Evolutionary Trends Continued 3. Feeding Adaptations • Suspension feeding of fixed organisms above the substrate, i.e. epifaunal suspension feeding, was the predominant type of feeding. • Various filtering devices evolved for filter-feeding: lophophores of bryozoans and brachiopods, tentacles of corals, gills of bivalves, brachioles and arms of crinoids and blastoids, and others. A. Invertebrate Evolutionary Trends Continued 3. Feeding Adaptations continued. • • • • • Abundant suspension feeders maximized their ability to obtain food in crowded competition by: • Feeding at a variety of heights above the sea floor, called tiering. Most food moves horizontally in bottom currents so tiered animals do not feed from the same water moving past them. • Invertebrates fed on a variety of food sizes. Bryozoans fed on particles <100 microns in size while cnidarians can eat small fish. Size selection of food was just another way of differentiating suspension feeder food resources. When the Ordovician began epifaunal suspension feeders were low in height with minimal tiering. With Ordovician invertebrate radiation came increased tiering with crinoids reaching 50 to 100 cm of height above the seafloor. First burrowing (shallow) infaunal suspension feeders accompanied increased predation. First significant predators (eurypterids and cephalopods). B. Rise and diversification. • First appearances occurred of corals, clams, bryozoans, starfish, echinoids or crinoids. • The articulate brachiopods were rare during the Cambrian but began a period of major diversification in the shallow-water marine environment. • Brachiopods dominated the Ordovician shallow water non-reef environments. • Types were very much facies related with inarticulates very restricted to finer sediments. • Most Ordovician brachiopods had a wide hinge and coarse ribs (e.g. Orthids). • Most Cambrian ones bore chitinous shells and were small. Ordovician brachiopods were larger and more diverse. 2 6/15/2015 B. Rise and diversification continued. • Foraminifera protozoans made their first appearance and increased in abundance and diversity later in the Paleozoic. • Ostracods first appeared and became “large” and abundant in some limestone. • The first corals and tabulate (always colonial) corals appeared in the Lower Ordovician but were small and subordinate to tabulate corals and stromatoporoids. • Crinoids became abundant for the first time. • Stalked echinoderms including crinoids and blastoids diversified. Although rare, the first crinoidal limestone (limestone with an abundance of crinoids) was deposited. B. Rise and diversification continued. • “Cystoids”, a heterogeneous group of blastozoans, reached their maximum abundance in the Ordovician and Silurian but were never very abundant. • Very early in the period primitive starfish and brittle stars appeared and in the Middle Ordovician the first sea urchin (echinoid) joinrd in. • Bryozoans first appeared and diversified greatly becoming a major contributor to Ordovician limestone. • Trepostomes (“stony”) were the dominant bryozoans. Many of these were branching stick-like or compact encrusting colonies. • Three other orders appeared. • In limestone and limy shale, bryozoans and brachiopods are the most common invertebrate components. B. Rise and diversification continued. • Bivalves first appeared near the base of the Middle Ordovician, radiated and became successful burrowing filter feeders but were not yet particularly abundant. • Gastropods diversified greatly. • An increased diversity and abundance of acritarchs (organic-walled phytoplankton of unknown affinity), the major phytoplankton group of the Paleozoic Era, occurred. Acritarchs were the primary food source of the suspension feeders that were becoming more common (more in Module 8). • Trilobites reached their greatest abundance and diversity early in the period. 3 6/15/2015 C. Declines in invertebrate groups and diversity. • Invertebrate grazers (gastropods) decimated the algal stromatolites that were so widespread before diversification of metazoans. • Late in the period trilobites began a long decline in abundance and diversity until their extinction shortly before the end of the Paleozoic. D. Dominant invertebrates. • Ordovician invertebrate communities dominated by articulate brachiopods, bryozoa, and corals. • Corals and bryozoa became primary reef builders. Other reef builders included stromatoporoids and other sponges. • Most of the benthic community was composed of bryozoans, brachiopods, corals, stalked echinoderms, sponges, bilves, gastropods, graptolites, trilobits, starfish, cephalopods, and conodonts. E. Lesser contributors to Ordovician assemblages. • Cystoids, echinoids, bivalves, gastropods, non-stromatoporoid sponges, ostracods, graptolites. 4 6/15/2015 F. Predators • Common large predators, including shelled cephalopods. • Rare and small in the Late Cambrian. • The cephalopods were the dominant and fastest predators of the Ordovician and were very common. • Only 1 order in Cambrian but 10 in Ordovician. • Most were open coiled (cyrtocone) or straight (orthocone). • Some got between 15 and 30 feet and were the largest animals yet to live. Eurypterids were significant predaceous arthropods. • Cuticle skeleton so most fossils are carbon films. • Most 10-20 cm but some up to 1.8 meters. G. Reefs. • Stromatoporoids • Became a major reef forming group in the upper Middle Ordovician. • Sponges, multilayered structure with small mounds on each layer and star shaped canals centered on mounds. • Interior usually recrystallized. • Major reef formers to end of Devonian. • Colonial rugose and tabulate corals began contributing to reefs in the Middle Ordovician G. Reefs. • Development of an Ordovician reef: • stabilization stage- stalked crinoids • colonization stage- stromatoporoids and bryozoans • diversification stage- corals, bryozoans, sponges, and stromatoporoids • domination stage- stromatoporoids 5 6/15/2015 H. Guide fossils • The three fossil groups most useful for Ordovician biostratigraphic correlation are the articulate brachiopods, graptolites, and conodonts. • Graptolites • Pelagic and benthic colonial related to modern hemichordates. • Along with the pelagic trilobites among the only fossils found in deep water shale. • 13 zones in European Ordovician. • Because some were pelagic they are very important in intercontinental correlation. • Most frequently found in organic shale. • Conodonts (see Module 8) • Very abundant. • Conodonts (phosphatic) are teeth of an eel-like creature. • Chitinozoa (see Module 8) • Extinct palynomorphs of unknown systematic position. • Brachiopods I. Vertebrates • Vertebrates arose from Hemichordates (partial notochords), which include modern acorn worms and extinct graptolites. • Oldest from Cambrian Burgess Shale (Pikaia) • Jawless vertebrates evolved in Late Cambrian, the armored Agnaths. • Filter feeders J. Plants. • First very rare evidence of land plants in the form of cuticle and spores. 6 6/15/2015 K. Extinction. • • • • End of the Ordovician marked by a mass extinction. 57% generic extinction. More than 100 families of marine invertebrates became extinct. Large-scale loss of diversity in many groups, particularly stalked crinoids, brachiopods, and trilobites. 7