The Great Ordovician Biodiversification Event

Critical Moments and Perspectives in Earth History and Paleobiology

Critical Moments and Perspectives in Earth History and Paleobiology

David J. Bottjer, Richard K. Bambach, and Hans-Dieter Sues, Editors

The Emergence of Animals: The Cambrian Breakthrough

Mark A. S. McMenamin and Dianna I. S. McMenamin

Phanerozoic Sea-Level Changes

Anthony Hallam

The Great Paleozoic Crisis: Life and Death in the Permian

Douglas H. Erwin

Tracing the History of Eukaryotic Cells:The Enigmatic Smile

Betsy Drexler and Robert Allan Obar

The Eocene-Oligocene Transition: Paradise Lost

Donald R. Prothero

The Late Devonian Mass Extinction: The Frasnian/Famennian Crisis

George R. McGhee Jr.

Dinosaur Extinction and the End of an Era: What the Fossils Say

J. David Archibald

One Long Experiment:Scale and Process in Earth History

Ronald E. Martin

Interpreting Pre-Quaternary Climate from the Geologic Record

Judith Totman Parrish

Theoretical Morphology:The Concept and Its Applications

George R. McGhee Jr.

Principles of Paleoclimatology

Thomas M. Cronin

The Ecology of the Cambrian Radiation

Andrey Yu. Zhuravlev and Robert Riding, eds.

Plants Invade the Land:Evolutionary and Environmental Perspectives

Patricia G. Gensel and Dianne Edwards, eds.

Exceptional Fossil Preservation: A Unique View on the Evolution of Marine Life

David J. Bottjer, Walter Etter, James W. Hagadorn, and Carol M. Tang, eds.

EDITED BY

Barry D. Webby, Florentin Paris, Mary L. Droser, and Ian G. Percival

Columbia University Press

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E-ISBN 978-0-231-50163-7

Library of Congress Cataloging-in-Publication Data

The great Ordovician biodiversification event / Barry D. Webby, Florentin Paris, Mary L. Droser and Ian G. Percival, editors.

p. cm.—(Critical moments and perspectives in Earth history and paleobiology)

Includes bibliographical references and index.

ISBN (cloth): 0231-12678-6

1. Biodiversity—Ordovician. 2. Paleontology—Ordovician. 3. Earth History—Ordovician. I. Webby, Barry D., 1934- II. International Geological Correlation Program, Project No. 410 (Ordovician biodiversity). III. Series

A Columbia University Press E-book.

CUP would be pleased to hear about your reading experience with this e-book at cup-ebook@columbia.edu.

This book is dedicated to all paleontologists who, over the past 200 years, have described the Ordovician biotas of the world. Their contributions have provided the knowledge base that our team of specialists have drawn on for this comprehensive stocktaking of arguably the greatest-ever interval of sustained diversification of life on earth.

Acknowledgments

1   Introduction

Barry D. Webby

PART I

Scaling of Ordovician Time and Measures for Assessing Biodiversity Change

2   Stratigraphic Framework and Time Slices

Barry D. Webby, Roger A. Cooper, Stig M. Bergström, and Florentin Paris

3   Calibration of the Ordovician Timescale

Peter M. Sadler and Roger A. Cooper

4   Measures of Diversity

Roger A. Cooper

PART II

Conspectus of the Ordovician World

5   Major Terranes in the Ordovician

L. Robin M. Cocks and Trond H. Torsvik

6   Isotopic Signatures

Graham A. Shields and Ján Veizer

7   Ordovician Oceans and Climate

Christopher R. Barnes

8   Was There an Ordovician Superplume Event?

Christopher R. Barnes

9   End Ordovician Glaciation

Patrick J. Brenchley

10   Ordovician Sea Level Changes: A Baltoscandian Perspective

Arne Thorshøj Nielsen

PART III

Taxonomic Groups

11   Radiolarians

Paula J. Noble and Taniel Danelian

12   Sponges

Marcelo G. Carrera and J. Keith Rigby

13   Stromatoporoids

Barry D. Webby

14   Conulariids

Heyo Van Iten and Zdenka Vyhlasová

15   Corals

Barry D. Webby, Robert J. Elias, Graham A. Young, Björn E. E. Neuman, and Dimitri Kaljo

16   Bryozoans

Paul D. Taylor and Andrej Ernst

17   Brachiopods

David A. T. Harper, L. Robin M. Cocks, Leonid E. Popov, Peter M. Sheehan, Michael G. Bassett, Paul Copper, Lars E. Holmer, Jisuo Jin, and Rong Jia-yu

18   Polyplacophoran and Symmetrical Univalve Mollusks

Lesley Cherns, David M. Rohr, and Jiří Frýda

19   Gastropods

Jiří Frýda and David M. Rohr

20   Bivalve and Rostroconch Mollusks

John C. W. Cope

21   Nautiloid Cephalopods

Robert C. Frey, Matilde S. Beresi, David H. Evans, Alan H. King, and Ian G. Percival

22   Tube-Shaped Incertae Sedis

John M. Malinky, Mark A. Wilson, Lars E. Holmer, and Hubert Lardeux

23   Worms, Wormlike and Sclerite-Bearing Taxa

Olle Hints, Mats Eriksson, Anette E. S. Högström, Petr Kraft, and Oliver Lehnert

24   Trilobites

Jonathan M. Adrain, Gregory D. Edgecombe, Richard A. Fortey, Øyvind Hammer, John R. Laurie, Timothy McCormick, Alan W. Owen, Beatriz G. Waisfeld, Barry D. Webby, Stephen R. Westrop, and Zhou Zhi-yi

25   Eurypterids, Phyllocarids, and Ostracodes

Simon J. Braddy, Victor P. Tollerton Jr., Patrick R. Racheboeuf, and Roger Schallreuter

26   Crinozoan, Blastozoan, Echinozoan, Asterozoan, and Homalozoan Echinoderms

James Sprinkle and Thomas E. Guensburg

27   Graptolites: Patterns of Diversity Across Paleolatitudes

Roger A. Cooper, Jörg Maletz, Lindsey Taylor, and Jan A. Zalasiewicz

28   Chitinozoans

Florentin Paris, Aïcha Achab, Esther Asselin, Chen Xiao-hong, Yngve Grahn, Jaak Nõlvak, Olga Obut, Joakim Samuelsson, Nikolai Sennikov, Marco Vecoli, Jacques Verniers, Wang Xiao-feng, and Theresa Winchester-Seeto

29   Conodonts: Lower to Middle Ordovician Record

Guillermo L. Albanesi and Stig M. Bergström

30   Vertebrates (Agnathans and Gnathostomes)

Susan Turner, Alain Blieck, and Godfrey S. Nowlan

31   Receptaculitids and Algae

Matthew H. Nitecki, Barry D. Webby, Nils Spjeldnaes, and Zhen Yong-yi

32   Acritarchs

Thomas Servais, Jun Li, Ludovic Stricanne, Marco Vecoli, and Reed Wicander

33   Miospores and the Emergence of Land Plants

Philippe Steemans and Charles H. Wellman

PART IV

Aspects of the Ordovician Radiation

34   The Ichnologic Record of the Ordovician Radiation

M. Gabriela Mángano and Mary L. Droser

35   The Ordovician Radiation: Toward a New Global Synthesis

Arnold I. Miller

List of Figures and Tables

References

List of Contributors

Index

This book derives directly from a significant part of the work of UNESCO-IUGS–supported IGCP Project no. 410: The Great Ordovician Biodiversification Event: Implication for Global Correlation and Resources (1997–2002). As the first globally directed IGCP program to highlight Ordovician rocks and fossils, IGCP 410 established (1) closely coordinated regional teams for data collection and analysis of biodiversity; (2) a wholly integrated and well-calibrated timescale for global and regional correlation, and (3), from 1998, a complementary clade team work program to assess global biodiversity patterns in time and space Also a Web-based relational database managed by Arnold Miller (University of Cincinnati) was available for input of relevant biotal data.

We acknowledge particularly UNESCO and IUGS and the members of the IGCP Scientific Board, who gave IGCP 410 six years of continuing encouragement and support. They, together with a number of regional organizations, helped us sponsor highly successful meetings each year in different parts of the world.

The following individuals and their organizations were especially supportive in arranging our globally focused meetings: Tatjana N Koren’ (A. P. Karpinsky All-Russian Geological Research Institute—VSEGEI, St. Petersburg, Russia), Duck K. Choi (Institute of Geological and Environmental Sciences, Seoul National University, Korea), Rong Jia-yu, Zhou Zhi-yi, and Chen Xu (Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Academia Sinica, China), Wang Xiao-feng (Yichang Institute of Geology and Mineral Resources, MGMR, China), Petr Kraft and Oldřich Fatka (Institute of Geology–Palaeontology, Faculty of Science, Charles University, Prague, Czech Republic), John Talent, Ruth Mawson, and Glen Brock (Centre of Ecostratigraphy and Palaeobiology, Macquarie University, North Ryde, Sydney, Australia), Mary Droser (Department of Earth Sciences, University of California at Riverside, United States), Evgeny Yolkin (Institute of Petroleum Geology, Russian Academy of Sciences, Novosibisk, Russia), and Minjin Chuulin (School of Geology, Mongolian Technical University, Ulaanbaatar, Mongolia).

We maintained important linkages in our cooperative work programs with the IUGS/ICS Subcommission on Ordovician Stratigraphy (Chairman Stan Finney) and with IGCP project no. 421—North Gondwana Mid-Palaeozoic Biodynamics (co-leaders Raymund Feist and John Talent). A former chairman of the Ordovician Subcommission, Reuben J. Ross Jr., encouraged the publication of a worldwide series of 12 Ordovician correlation charts between 1980 and 1995 (IUGS Publications nos. 1–2, 6, 8, 11–12, 21–22, 26, 28–29, 31), and these were invaluable in the work to establish precise stratigraphic ranges for many of the taxa used in this survey.

From 1998 it was planned that our clade team work would include the compilation of an edited volume covering the biodiversity profiles of as many taxonomic groups as possible. The main clade team meeting of IGCP 410 was held at the University of California (Riverside) in June 2001. Organized by Mary Droser and her Riverside colleagues, it focused on a wide range of global and regional biodiversity topics across the main Ordovician taxonomic groups and the Ordovician timescale. Many chapter authors attended this most productive meeting, but a number of active, mainly overseas participants were unable to attend. All of these have been encouraged to submit contributions to the volume. In addition, a few other specialist contributors were solicited after the meeting to fill obvious gaps so that a wider global coverage could be presented in the book.

During the preparation of this book, Barry Webby benefited from being able to use the facilities of the Centre of Ecostratigraphy and Palaeobiology, the Department of Earth and Planetary Sciences, and the University Library of Macquarie University in Sydney.

In terms of the editorial processing, Alan Owen (trilobites) and Dave Harper (brachiopods) assisted by coordinating their large chapters. Roger Cooper’s advice and his contribution on diversity measures are also much appreciated. Collective thanks are extended, also, to all the other authors who contributed manuscripts to the volume; many, additionally, helped by critically reading drafts submitted by other authors in the project.

The editorial process could not have been undertaken in the available time frame without the substantial support of numerous referees worldwide. Special thanks to them all for their insightful reviews. They include Jonathan Aitchison, Frits Akterberg, Bob Anstey, Claude Babin, Arthur Boucot, Simon Conway Morris, Kent Condie, James Crampton, Dave Elliott, Ray Ethington, Dianne Edwards, Stan Finney, Dan Fisher, Michael Foote, Dan Goldman, Roland Goldring, Joe Hannibal, Charles Holland, Steven Holland, Richard Hoare, Warren Huff, Nigel Hughes, Peter Jell, Ron Johns, Anita Löfgren, Merrell Miller, Heldur Nestor, Paddy Orr, Robert Owens, Mark Patzkowsky, Geoffrey Playford, John Pojeta Jr., John Pickett, John Peel, June Philips Ross, Charles Ross, Enrico Serpagli, Paul Selden, John Shergold, Colin Stearn, Desmond Strusz, Julie Trotter, Fons VandenBerg, Jean Vannier, Pat Wilde, Henry Williams, A. (Tony) D. Wright, Gavin Young, Ellis Yochelson, and three reviewers who wish to retain their anonymity.

The historical scientist focuses on detailed particulars—one funny thing after another—because their coordination and comparison permits us, by consilience of induction, to explain the past with… confidence (if the evidence is good).

STEPHEN JAY GOULD, Wonderful Life

This introductory chapter presents the scope, aims, and organization of the volume; outlines previous work on Ordovician biodiversity topics; and gives an overview of the chapters in the volume, from those briefly appraising the Ordovician World to those more comprehensively surveying the diversification patterns of the main Ordovician taxonomic groups. The chapter ends with some closing remarks on the Ordovician Radiation and future directions.

Scope and Aims

Two of the greatest evolutionary events in the history of life on earth occurred during Early Paleozoic time. The first was the Cambrian explosion of skeletonized marine animals—what Wilson (1992: 188) called the big bang of animal evolution—about 540 million years (m.y.) ago. The second was the Great Ordovician Biodiversification Event, the focus of this book. During the 46-m.y. span of Ordovician time (489–443 m.y. ago), an extraordinarily varied range of evolutionary radiations of Cambrian-, Paleozoic- and Modern-type biotas appeared, the most diversified occurring in marine continental platform to open ocean habitats. Animals (arthropods) also first walked on land, and based on their cryptospore record, the first nonvascular bryophytelike plants colonized damp areas on land.

The Cambrian explosion of skeletonized animals is now comparatively well documented, though the timing of the initial event still needs to be reassessed (Bowring and Erwin 1998). Books have been published on a number of aspects of this explosion, for example, Glaessner’s The Dawn of Animal Life: A Biohistorical Study (1984), Lipps and Signor’s Origin and Early Evolution of the Metazoa (1992), and Zhuravlev and Riding’s The Ecology of the Cambrian Radiation (2000). Also well documented is the history of the extraordinary Mid Cambrian Burgess Shale faunas, about 505 m.y. ago. The most notable publications are Gould’s Wonderful Life (1989), Briggs, Erwin, and Collier’s The Fossils of the Burgess Shale (1994), and Conway Morris’s The Crucible of Creation (1998b).

In contrast, until very recently, there was no Ordovician biodiversity volume, or at least none that focused significantly on aspects of Ordovician biodiversity. However, nearly half the contributions in Crame and Owen’s Palaeobiogeography and Biodiversity Change: The Ordovician and Mesozoic-Cenozoic Radiations (2002) have addressed issues relating to Ordovician diversity change. The important role of plate tectonics during the Ordovician was singled out—examples, such as the fragmentation of the Gondwanan margin, the drift of Avalonia, and the development of an array of marginal to oceanic terranes (including island chains) that effectively partitioned ocean circulation patterns within the Iapetus Ocean—as prominent in engendering the patterns of diversity change.

Ordovician biodiversity has typically been studied in three different ways: (1) taxonomic diversity, which involves a focus on the taxonomic richness and turnover (originations and extinctions) within fossil groups; (2) ecologic diversity, which examines how organisms (or their communities) adapt to fill niche spaces in order to exploit available food resources more successfully; and (3) morphological diversity (usually termed disparity), which traces the patterns of morphological (design) change in various fossil groups.

This book is devoted primarily to documenting the taxonomic diversification of Ordovician biotas, in both global and regional contexts, and to firming up the timing of the most important diversification events. The volume has a broad coverage, but inevitably there are gaps. A few aspects of ecologic diversity have been treated, but the coverage is limited, and aspects of morphological diversity (disparity) are barely touched on. Moreover, only limited discussion of possible causes for the radiation events is included.

All these Ordovician diversification events form part of the Great Ordovician Diversification Event, also called the Ordovician Radiation (Droser et al. 1996). Although the most intensive part of the Ordovician Radiation was during the Mid to Late Ordovician epochs, an interval of 28 m.y. (until the second extinction pulse of the end Ordovician mass extinctions—Sheehan 2001b; chapter 9 in this volume), some taxonomic groups (e.g., trilobites, inarticulated brachiopods, graptolites, conodonts, and rostroconch mollusks) also diversified significantly during the Early Ordovician. Consequently, all these evolutionary events from the beginning to virtually the end of the Ordovician Period—through nearly 46 m.y. of earth history—should be treated as belonging to the Ordovician Radiation (figure 1.1).

FIGURE 1.1. (A) Phanerozoic taxonomic diversity of marine animal families (slightly modified from Sepkoski 1984: figure 1). The fields Cm, Pz, and Md represent the Cambrian, Paleozoic, and Modern evolutionary faunas (EFs), respectively. Note also the generalized field of additional, poorly preserved families at the top of the diagram. Sepkoski’s timescale follows Harland et al. (1982) and includes the following abbreviations: V = Vendian; = Cambrian; O = Ordovician; S = Silurian; D = Devonian; C = Carboniferous; P = Permian; TR = Triassic; J = Jurassic; K = Cretaceous; T = Tertiary. (B) The Middle to Upper Cambrian, Ordovician, and Silurian (except Pridoli) taxonomic diversity of marine animal genera (modified from Sepkoski 1995: figure 1). The main field of Cambrian, Paleozoic, and Modern EFs is shown, as well as Sepkoski’s time units, including British series for his Ordovician and Silurian subdivisions. His abbreviations are as follows: M = Middle Cambrian; U = Upper Cambrian; T = Tremadocian; Ar = Arenig; Ln = Llanvirn; L = Llandeilo; C = Caradoc; As = Ashgill; Ly = Llandovery; W = Wenlock; Lv = Ludlow. The Llandeilo series has now been abandoned in favor of an enlarged Llanvirn, the name Llandeilian retained for its upper stage (figure 2.1), and the overlying Caradoc expanded downward to fill the gap (Fortey et al. 1995, 2000). Note also the added scale bar at the bottom of the diagram that comprises the tripartite global series subdivisions for the Ordovician System, radiometric dates in millions of years (Ma), and the time-slice subdivisions used in this volume.

No comparable, well-defined major extinction episode has been recognized at the beginning of the Ordovician Period. According to Sepkoski (1981b, 1995, 1997), after the dramatic Early Cambrian radiations there was a phase of apparent stagnation or quiescence that persisted through the Mid Cambrian to the Early Ordovician, with a comparatively lower overall diversity thoughout (figure 1.1B). Yet it remained an interval of high turnover—with high extinction rates limiting the overall buildup of diversity, including the well-defined successive pulses of mainly trilobite-based Mid to latest Cambrian biomere extinctions (Palmer 1979; Ludvigsen 1982; Zhuravlev 2000). The last of these more or less coincides with the base of the North American Ibexian (Ross et al. 1997), that is, three conodont zones below the base of the Ordovician (Cooper et al. 2001). A few extinction horizons have been identified within the Ordovician, but they all appear to be relatively minor pulses that have yet to be demonstrated as being truly global events. For example, Ji and Barnes (1993, 1996) documented an Early Ordovician (mid Tremadocian) conodont extinction event in Laurentia; Sepkoski (1992b, 1995, 1996) reported extinction horizons in the late Mid Ordovician (near the bottom and top of the Darriwilian, respectively); and Patzkowsky et al. (1997) identified a climatic event with associated Late Ordovician (mid Caradoc) brachiopod-dominated extinction in Laurentia. The last has wider, probably even global, importance, correlating with climatic and other changes—including the late Keila extinctions of chitinozoans, acritarchs, and ostracodes—in Baltoscandia (Kaljo et al. 1996; Ainsaar et al. 1999).

Biotas

This book was compiled by a large team of Ordovician specialists from around the world, under the aegis of the joint UNESCO and International Union of Geological Sciences (IUGS) geosciences initiative—the International Geological Correlation Programme (IGCP)—which supported IGCP project no. 410. The aim in establishing this first globally oriented, internationally sponsored IGCP project was to have the Great Ordovician Biodiversification Event comprehensively evaluated in a collective effort. When IGCP 410 became formally established in 1997, data collection and analysis of biotas were coordinated mainly on a regional basis, but in 1998 a separate study program was developed with a more constrained global approach on the major fossil groups. This became an evaluation of how each taxonomic group diversified through Ordovician time, with assessments of patterns of diversity change, and rates of origination and extinction, based on the assembly of genus- and species-level taxonomic diversity data of the various fossil groups. Leading specialists and their colleagues were invited to participate in the three-year work program and to attend a major meeting organized by Mary Droser in June 2001 at the University of California in Riverside to discuss the results of the compilations. The book derives in part from these contributions, with a number of additional studies from other specialists added to the project after the Riverside meeting to widen the coverage to include nearly all fossil groups.

The leader of each fossil group chose his or her own team of co-workers to tackle the compilation of data and assembly of the manuscript for the particular chapter. The groups varied in size from large (with a coordinating author and many coauthors) to small, single-author presentations. We encouraged each team to establish its diversity surveys so that they highlighted patterns of diversity change, originations, and extinctions, where possible down to species level at least for the pelagic groups, and to genus, and where possible to species level for benthic groups. Most authors assembled their primary taxonomic data using their own databases (spreadsheets or census lists).

The pelagic groups (e.g., graptolites, chitinozoans, radiolarians) and some benthic groups (e.g., bryozoans, sponges, stromatoporoids, echinoderms) are presented as species-level, or combined species- and genus-level, diversity surveys. Larger benthic groups (e.g., trilobites, brachiopods, gastropods, bivalves, nautiloids) have been surveyed mainly at the genus level at this stage. Although we attempted to provide the widest possible coverage of Ordovician biotas, it was inevitable that some unevenness in the levels of documentation and analysis would occur from chapter to chapter. For a variety of reasons, a number of groups are presented with rather incomplete global diversity analyses. This may be because (1) only part of the group’s Ordovician record has been treated; (2) reliable data of well-preserved and diagnostic material are available only from one or two regions in the world, so the survey does not have a global focus; (3) genus- and species-level assignments of a group are unreliable or at least need further revision before a worldwide analysis can be attempted; or (4) present-day expertise on a particular group is lacking. Consequently, for a few groups, only preliminary statements could be included, and in some cases these assessments remain predominantly regionally based.

Although we have attempted to include most Ordovician groups in this genus- and species-level coverage (table 1.1), there are some gaps in the documentation—for instance, the Mid to Late Ordovician conodont diversity record and a few small groups of comparatively limited Ordovician occurrence, for example, foraminiferans (Lipps 1992a, 1992b), hydroids (Stanley 1986; Foster et al. 1999), and hyolithelminths. In addition, a few groups of microorganisms—such as two unicellular planktic green algal prasinophyte groups (leiospheres, tasmanitids) and the prolific, organic-walled, benthic cyanobacterium Gloeocapsomorpha prisca, a microorganism of considerable economic interest because it forms significant matlike accumulations of late Mid to Late Ordovician kukersite oil shales in the intracratonic basins of Estonia (Körts 1992), North America (Jacobson et al. 1988), and Australia (Foster et al. 1990)—have not been included in the survey.

An estimate of total numbers of Ordovician genera and species for each fossil group treated in the book (table 1.1) is provided by authors for general guidance only. It is not intended as a comprehensive listing of totals for all Ordovician groups. The list comprises about 4,605 genera (excluding the trace fossils). The total number of animal genera (less the plants—algae, acritarchs, miospores) is 4,254 genera. Earlier, Sepkoski (1995) employed a database of 4,367 animal genera (i.e., 12 percent of all known animal genera in the Phanerozoic) to prepare his outline of Ordovician diversity history. Only the few small groups mentioned earlier, the late Mid to Late Ordovician conodont record and about 70 genera of nautiloids from China and Russia, have been excluded from our present survey.

With recent publication of Sepkoski’s (2002) comprehensive listing of all known fossil marine animal genera, it is now possible to make more-up-to-date comparisons between the two lists of Ordovician animal generic diversity records (table 1.2). However, it is difficult to compare the two lists in detail because they represent completely different types of fossil data assemblies. The present study was not intended to be comprehensive but rather to involve as many available Ordovician specialists as possible in a stocktaking of their fossil group. In some cases these workers have adopted revised taxonomic categories from those listed in Sepkoski’s Compendium of Fossil Marine Animal Genera (2002). The Compendium represented a mammoth compilation of Phanerozoic animal genera, assembled from the data coverage of animal groups in Moore et al., Treatise on Invertebrate Paleontology (1953–1992), and many hundreds of other literature sources published up to 1998. Nevertheless, there is a remarkable degree of similarity between the two lists. The comprehensive Sepkoski (2002) compilation also provides a means to appreciate the full range of Ordovician taxonomic groups, including those biodiversity records that have not been documented here.

TABLE 1.1. A Preliminary Listing of Genus- and Species-Level Totals for the Ordovician Fossil Groups

Source: Compiled from data supplied by individual authors.

Note: This compilaton of animals, plants, and trace fossils is provided for general guidance only because some genus-level and especially the species-level taxonomy of fossil groups remains in a state of flux. For many groups a large backlog of taxonomic revision work still needs to be done.

Bivalve and scolecodont species data are shown as ranges, namely, between 530 and 680 and between 500 and 1,000 species, respectively. For entries of nautiloid genera and conulariid species, two numbers are included (one in parentheses): the lower number is the actual count (a mainly regional summation), and the higher number in parentheses is the global estimate. For the phyllocarids, the valid genera and species numbers are shown, plus the numbers of doubtful taxa (with a question mark). For trace fossils, only numbers of form genera (ichnogenera) could be supplied.

Abbreviations: c. = about; n.d. = data not determined (or not supplied).

TABLE 1.2. Comparative Generic Lists of Ordovician Animal Diversity Data

Source: Compiled from data supplied by individual authors (this study) and from Sepkoski (2002).

a Stromatoporoids are here listed (in this study and in Sepkoski’s list) as separate from other sponges.

b The problematic receptaculitids is the only group omitted from Sepkoski’s list. They are included in this study as a possible metazoan group.

c The bellerophontids have been included with the symmetrical univalves of this study, whereas they are included in Sepkoski’s list with the gastropods.

d The genus (Sphenothallus) is included in Sepkoski’s list as a conulariid.

e The genus (Byronia) is included in Sepkoski’s lists as a hyolithelminth.

Only a comparatively few groups have good-quality assignments of species data (table 1.1). These include mainly pelagic groups such as the graptolites, conodonts, chitinozoans, and radiolarians, but correct species assignments have also been established for a few benthic groups (e.g., stromatoporoids, conulariids, bryozoans, rostroconchs, eurypterids, echinoderms, and receptaculitids).

Wilson (1992) has an estimate of 1.4 million organisms (plants, animals, and microorganisms) currently known to be living on earth, though millions of insects, microbes, and other organisms remain undescribed across habitats from rain forests to the ocean deeps (Wilson 1992; Thorne-Miller 1999). The problems of estimating fossil species numbers is a more daunting task. Paul (1998) reviewed the available approaches, namely, to establish what proportion of living animal species is likely to be preserved in modern settings, as well as to establish the relationships between soft-bodied and skeletonized organisms in fossil Lagerstättern such as the Middle Cambrian Burgess Shale (Conway Morris 1986). These two approaches produce rather similar results, suggesting to Paul that only about 10 percent of all Phanerozoic species are likely to have been preserved. In assessing the Ordovician global record, we currently have only reasonably complete numbers for the genera—some 4,600 known Ordovician genera. From the generic and specific data in the 26 listed fossil groups (table 1.1), there are on average about five species for each genus. Multiplying this value by the generic total gives the very approximate total of 23,000 Ordovician skeletonized species. If this estimate represents only 10 percent of species likely to be preserved (Paul 1998), then the overall total (including the soft-bodied organisms) may be of the order of 200,000 to 250,000 Ordovician species.

Ordovician Time and Time Slices

A specific aim of IGCP 410 was to develop a more highly integrated, well-calibrated Ordovician timescale capable of providing a more reliable global and regional basis for correlating range data and establishing age relationships of the diverse biotas. It has long been recognized (Jaanusson 1960, 1979) that establishing biostratigraphic correlations using Ordovician biotas, especially on a global scale, is a difficult task because of the extensive biogeographic and ecologic differentiation of the faunas. Furthermore, the comparative lack of good reliable radiometric ages has limited the degree to which age relationships could be determined (Webby 1998). The priority to establish a stabilized global stratigraphic framework for the Ordovician System has been the main responsibility of the International Subcommission on Ordovician Stratigraphy (ISOS). It is an independent subcommission of the International Commission on Stratigraphy (ICS), and both ISOS and ICS exist under the umbrella of IUGS. Since the mid-1980s, ISOS has made good progress toward the establishment of a single set of globally based Series and Stage divisions, using the most highly resolved biologic indices (graptolite, conodont, and chitinozoan zones) coupled with available physical and chemical tools (table 1.3).

Stratigraphers, especially those closely associated with global correlation work of international stratigraphic bodies like ICS and ISOS, have long been aware of the need to draw clear distinctions between chronostratigraphic units based on rock sequences and geochronologic units that reflect intervals of geologic time (Salvador 1994). The terminology is straightforward for the bulk of conventional hierarchical usages, such as system, series, and stage with their geochronologic counterparts period, epoch, and age. However, most series have tripartite subdivisions, with names derived from their position in the system, for example, Lower Ordovician Series, Middle Ordovician Series, and Upper Ordovician Series. The nomenclature Early Ordovician Epoch and Late Ordovician Epoch is also acceptable, but the terminology for the intervening epoch has remained a problem because the same name Middle Ordovician has been applied to both series (for the rocks) and epoch (for time) usages. Only British stratigraphers (see Holland et al. 1978) have drawn a distinction in their regional work between middle (for rocks) and mid- (for time). According to S. C. Finney, who is the current chair of ISOS and second vice-chair of ICS, the terms Middle Ordovician and Mid Ordovician are now used for the series and its correlative epoch, respectively (Finney et al. 2003:351). Following such recent moves at the international level, we have adopted the formalized global names Middle Ordovician Series for the rock sequences and Mid Ordovician Epoch for time throughout this book. The categories of lower, middle, upper (for rocks) and early, mid, late (for time) have similarly been recognized for ratified, tripartite, global stage subdivisions and for informal, mainly lower-rank, usages, as depicted in table 1.3.

Of critical importance, especially for biodiversity studies, was the level of resolution of the timescale (Erwin 1993; Conway Morris 1998a). Clearly it was essential to establish the most refined timescale with the shortest-term subdivisions for the widest applicability in studies of global biodiversity change. Consequently, two new, highly integrated and resolved correlation charts were prepared by a small group of specialists in IGCP 410, in cooperation with members of ISOS. These present the global and regional stratigraphic frameworks, main biostratigraphic (graptolite, conodont, and chitinozoan) schemes for establishing cross ties, key radiometric dates, and a set of 19 time-slice divisions by Webby et al. (see chapter 2: figures 2.1, 2.2).

Sadler and Cooper’s (chapter 3: figure 3.1) new calibration of the Ordovician timescale based on computer-assisted constrained optimization (CONOP) procedures allowed the subdivisions in the correlation charts to be calibrated with respect to geologic time. CONOP calibrations involved processing a huge data set of first- and last-appearance events based on 1,100 or so Ordovician and Silurian graptolite species from nearly 200 measured sections. A best fit sequence of events was derived in the CONOP process to establish a relative timescale from the thousands of event levels. The scale became numerically calibrated when radiometric dates and ranges of key graptolites (and conodonts) were interpolated along a regression line.

TABLE 1.3. Summary of the Global Subdivisions of the Ordovician System/Period

Ratified global subdivisions are shown in bold and are capitalized; informal stage/age and substage/subage usages are not capitalized; estimated radiometric ages (Ma) for the base and top of the Ordovician are shown in square brackets; N/A = subdivisions not applicable.

Note that a distinction is drawn between the System, or body of rocks, and the Period representing geologic time. The 19 time slices adopted for use in this book are shown in the right column.

a The British Hirnantian stage is equivalent to the upper/late Ashgill interval.

b The British Llanvirn is correlative with the middle/mid to upper/late parts of the Darriwilian Stage.

c The British lower/early Arenig and the middle/mid Arenig are more or less correlative with the unnamed upper Lower/late Early Ordovician and lower Middle/early Mid Ordovician global stages/ages, respectively.

In a more recent development, Cooper and Sadler (2002) have shown that the CONOP application can be used directly, without the time unit bias, to establish a global diversity curve. Again taking the well-studied and widely distributed graptolites, they have depicted a running diversity curve of 2,272 estimates of interval free graptolite species standing diversity through Ordovician and Silurian time.

The CONOP calibrations (chapter 3) for this biodiversity project have added significantly to establishing the Ordovician timescale as an integrated, high-resolution dating and correlation tool, to a point where it is now arguably one of the best-resolved intervals of time within the Paleozoic Era. This indicates, despite Jaanusson’s (1960, 1979) earlier misgivings, that the extreme levels of biogeographic and ecologic differentiation of the Ordovician biotas were not limiting factors. The 19 slices employed for the Ordovician Period (46 m.y. duration; table 1.2), with time-slice intervals of relatively equal units of time (intervals that each span between 1.6 and 3.0 m.y.), provide a most practical, best-resolved, and reliable basis for use in this present biodiversity survey. These standardized time slices, as emphasized by Cooper (1999c:441), are as fine as is practical to allow precision in global studies of diversity change of the fossil groups. Two types of abbreviations are employed through this work to distinguish between time, as a duration or interval in millions of years (m.y.), and time as a specific age assignment including radiometric ages in millions of years (Ma). Authors of the chapters in this volume were provided with the correlation charts (chapter 2: figures 2.1, 2.2) so that time equivalences could be maintained in data compilations across the different groups. In addition, abbreviated left- and bottom-margin versions of the key elements of these Ordovician stratigraphic charts were supplied so that they could be incorporated at the margins of diversity plots when compiling diversity data and printed with a consistent time frame for comparative purposes throughout the chapters of the book. The majority of authors have, in compiling their diversity data, successfully employed the standardized global time-slice scheme across a wide range of environmentally and provincially differentiated profiles.

Diversity Measures

Another important aspect of this biodiversity study was to have some consistency in the kinds of measures used in the volume for summarizing the data on diversity patterns and turnover rates. A short chapter on diversity measures is contributed by Cooper (chapter 4). The guidelines distributed to contributors provided general recommendations for establishing the best estimate of the true mean standing diversity, irrespective of the length of the time unit and taxonomic rank (Cooper 1999c), and the means of determining origination and extinction rates, the best approach being to estimate the rates in million-year intervals, which, of course, is dependent on the reliability of the calibration of the timescale (see earlier in this chapter). The measures have been adopted by about half the contributors, giving overall a relatively uniform set of presentations without limiting flexibility for those authors who wished to use other measures to emphasize particular features in their databases. Some additional measures that were circulated in a final update of the guidelines could not be adopted by some authors because they had already compiled their data and submitted their manuscripts.

Cooper’s standardized symbols (d, o, e, and subscript i, for diversity, originations, extinctions, and interval in m.y.) have been used widely throughout the book. On the other hand, no symbols were recommended to denote the taxonomic level of the compilation (genus or species), as it was thought preferable to explain the hierarchical level of the compilation in the text or in an appropriate caption. Cooper’s (chapter 4) contribution is important in identifying normalized diversity (dnorm) as consistently providing the best estimate of true mean standing diversity; hence it is the most appropriate measure for determining diversity change through geologic time.

With the highly resolved and calibrated global and regional time frame, the contributors of the various fossil groups were able to plot their data (genera and/ or species) accurately as global and/or regional stratigraphic ranges and then, using the diversity measures provided, derive the patterns of diversity change and turnover rates from their plotted range-chart data.

The diversity data have been, in some cases, plotted in two sets of diversity curves, one based on actual sampled diversity through time slices and the other as range-through diversity, whereby a taxon is shown as having a continuous range, though absent from a time slice (possibly owing to sampling failure or biofacies shift) between occurences in the immediately preceding and succeeding time slices (for examples, see Regional Patterns in Australia and New Zealand, the Anglo-Welsh Sector of Avalonia and in South China, chapter 24).

Organization

The present book derives from the contributions of 96 Ordovician paleontologists, stratigraphers, geochemists, and other geologists from 17 countries throughout the world. The book comprises this introduction and thirty-four other chapters, which have been divided into four main sections: part I: Scaling of Ordovician Time and Measures for Assessing Biodiversity Change; part II: Conspectus of the Ordovician World; part III: Taxonomic Groups; and part IV: Aspects of the Ordovician Radiation.

The three chapters of part I deal with the stratigraphic framework, time slices, and calibration of the Ordovician timescale (chapters 2 and 3) and the measures of diversity (chapter 4) that formed the basis of this biodiversity study. Outlines for these chapters have been presented earlier.

The remainder of this introductory chapter comprises (1) a brief history of earlier work on Ordovician biodiversity topics and (2) an overview of all the remaining chapters, divided into two sections, for part II (chapters 5–10) under the subheading Ordovician World in Brief and for parts III and IV (chapters 11–35) under the subheading Synopses of Fossil Groups.

Part II (Conspectus of the Ordovician World; chapters 5–10) includes a broad overview of a number of significant physical and chemical features of the Ordovician world, presented in order to (1) illustrate what marked contrasts exist between the Ordovician and the present-day climates and in the dispositions of continents and oceans and (2) provide background to some of the features that may have been influential in shaping this greatest diversification of marine life.

Part III (Taxonomic Groups; chapters 11–33) comprises a nearly complete coverage of taxonomic groups. For reasons discussed earlier in this introduction, the chapters vary considerably, from comprehensive, integrated surveys of global and regional genus- and/or species-level diversity data to far more preliminary compilations, focusing mainly on a few aspects of regional genus- or species-level biodiversity. The two concluding chapters of part IV (Aspects of the Ordovician Radiation; chapters 34 and 35) deal with the ichnofossil record and a global synthesis of the Ordovician Radiation.

Ordovician Biodiversity Perspectives: Earlier Work

Evolutionary Faunas (EFs) and Floras: Some Global Considerations

Faunal Patterns

Sepkoski’s (1978, 1979, 1981a, 1984, 1991a, 1997) numerous contributions on the patterns of the Phanerozoic global marine animal diversity change have given much attention to the development of the three Great Evolutionary Faunas (Cambrian, Paleozoic, and Modern EFs) and the pivotal role played by the Ordovician Radiation. The representatives of these EFs were recognized by Sepkoski as somewhat fuzzy sets of unrelated higher taxa with similar histories of diversification through extended intervals of time. At each level of taxonomic hierarchy (order, family, or genus), there were marked differences in the relative intensities of the Cambrian and Ordovician radiations. In comparison with the Cambrian increases, approximately twice as much ordinal biodiversity was added to the Ordovician marine system, some three times more familial diversity (figure 1.1A), and nearly four times more genus-level diversity (Sepkoski 1978, 1988; Sepkoski and Sheehan 1983). Each successive EF exhibited a greater diversity and ecologic complexity (Sheehan 2001a).

The Cambrian EF was an explosive evolutionary event involving appearances of the first well-skeletonized metazoans—groups such as trilobites, inarticulated brachiopods, hyoliths, monoplacophorans, and eocrinoids. The fauna expanded exponentially during the Early Cambrian, an interval of 20 m.y. (Sepkoski 1979, 1997: figure 1; Zhuravlev 2000); leveled off during the Mid to Late Cambrian, at a peak diversity of about 100 families; and then declined gradually through the Ordovician as the Paleozoic EF expanded. The Cambrian EF was reduced still further to about 30 families by the short-lived, glacially induced end Ordovician mass extinction. The fauna included appearances of a larger number of higher-level categories—most of the skeletonized metazoan phyla and nearly two-thirds of the classes (Sepkoski 1981a). On the other hand, lower-level categories were sometimes depauperate—rather plain, even grubby species, according to Valentine (1973: 451). The communities were mainly benthic, rather generalized, and intergrading, composed of surface deposit feeders, grazers, carnivores, and suspension feeders with low epifaunal or infaunal tiering. Overall the fauna exploited little of the potential ecospace (Bambach 1983).

This contrasts markedly with the patterns of global diversity change during the Ordovician Period. The main contributors in this new wave of major diversification were elements of Sepkoski’s (1979, 1984) Paleozoic EF. The fauna included the articulated brachiopods, cephalopods, crinoids, ostracodes, stenolaemate bryozoans, and corals. Components of the Paleozoic EF diversified very slowly in the Cambrian, with only slight increase during the Late Cambrian. In the Ordovician an exponential rise of diversity occurred, which was at a more rapid rate than at any other time during the Phanerozoic (Sepkoski 1995). The communities of attached epifaunal suspension feeders, deep burrowers, and carnivores greatly expanded (Bambach 1983; Droser and Sheehan 1997a), as did more specialized reef and hardground communities. Compared with those of the Cambrian EF, the taxa of the Paleozoic EF became more specialized, with the development of narrower ecologic requirements, use of resources, and competitive abilities. Suspension feeders of all types began to appear, along with the scavengers and carnivores that preyed on them from the earliest Ordovician (Signor and Vermeij 1994).

More than 350 new Paleozoic families were added in what Sepkoski (1981b:204) called the largest turnover in composition of marine faunas seen in the history of the oceans (figure 1.1A). The familial diversity curve, then, exhibits a sudden flattening after its steep upward climb, and this is interpreted to mean that the ecospace had finally been filled to capacity, to its equilibrium level, as shown by the nearly horizontal, Paleozoic-wide diversity plateau (Sheehan 2001a). However, two extinction/rebound perturbations disrupted the continuity of this diversity plateau; the first was the end Ordovician mass extinction, and the second was represented by the pulses of Late Devonian extinction. Despite these, the Paleozoic EF remained dominant for nearly 240 m.y. until it was far more severely disrupted in the end Permian mass extinction (Erwin 1993).

Zhuravlev (2000) noted that, with the exception of the bryozoans, all the phyla that participated actively in the major Ordovician diversifications were present in the Cambrian (e.g., precursors to chordates, conodonts and graptolites, the trilobites, brachiopods, sponges, echinoderms, and the mollusk groups, such as rostroconchs, cephalopods, gastropods, and polyplacophorans). Most of these groups (apart from trilobites and inarticulated brachiopods and, to a lesser extent, sponges and echinoderms) remained unimportant in Cambrian communities, but they gradually emerged from the shadows as opportunities arose following the repeated extinctions of late Early, Mid, and Late Cambrian time (Zhuravlev 2000). By the latest Cambrian (Sunwaptan) they had already taken over, in the sense that the last diversity peak was produced by them.

Sepkoski’s (1979, 1984) Modern EF had a comparatively limited development in the Ordovician, though its two dominant mollusk members (gastropods and bivalves) did expand markedly during Ordovician time (Sepkoski and Sheehan 1983; Sepkoski 1991a). Crustaceans, gymnolaemate bryozoans, foraminiferans, echinoids, fishes, reptiles, and mammals were other typical Modern EF components. The fauna exhibited a greater variety of predators and more highly complex infaunal communities (Bambach 1983, 1985; Thayer 1983). Such changes allowed ecospace to be divided more finely and still higher diversities to be achieved. The changes through the Ordovician Period mainly involved a very slow and steady increase in importance of members of the Modern EF (mainly bivalves and gastropods) relative to the overwhelmingly dominant members of the Paleozoic EF. This pattern continued through the Paleozoic (figure 1.1A), until the end Permian mass extinction, when differential survival of the faunal components occurred—the Paleozoic EF losing 79 percent of its familial diversity, compared with the Modern EF’s 27 percent loss (Sepkoski 1984). The Modern EF became the dominant fauna, and the expansion continues to the present with no sign of the appearance of a diversity plateau (Sepkoski 1997).

Overview of the Ordovician Diversity Record

Sepkoski (1995) presented a very important summary of the diversification history of Ordovician biotas that focused not only on all the marine animal genera (see figure 1.1B) but also on typical representatives of individual fossil groups, with separate diversity curves for each group. Figure 1.1B is basically a reproduction of Sepkoski’s original (1995) figure 1—that is, it includes his primary data compiled in Ordovician generic-level diversity curves of marine animal genera against 12 Ordovician time units derived from British series subdivisions, with a scale bar added at the bottom of his diagram, for comparative purposes, to illustrate relevant new global Lower, Middle, and Upper Ordovician series, radiometric dates, and time-slice data as used throughout this volume. Sepkoski (1995) established his 12 time unit subdivisions by dividing each British series into two, except for the Llandeilo, which he left undivided, and the Caradoc, which he divided into three. In addition, his lower Arenig and upper Ashgill subdivisions were allocated longer and shorter durations, respectively, in comparison with the other time units. Similar stratigraphic subdivisions have been used in Sepkoski’s (1992a, 2002) family- and generic-level compendia. Stratigraphic relationships between Sepkoski’s subdivisions and the global series and time-slice (TS) subdivisions used in this volume are shown in figure 1.1B, in particular, the base of the Middle Ordovician with a position toward the middle Arenig, and the base of the Upper Ordovician with a level close to the middle Llandeilo.

Sepkoski’s (1995) figure 1 (also reproduced here in figure 1.1B) for all the marine animal genera illustrates that the global diversity levels during the Early Ordovician were maintained at levels broadly similar to those of the preceding late Mid to Late Cambrian. Both of these Cambrian and Early Ordovician intervals were interpreted by Sepkoski (1995) as intervals of high turnover (i.e., including high rates of extinction, as discussed earlier) with several Paleozoic EF groups expanding slowly at the expense of Cambrian EF members through Late Cambrian to Early Ordovician time. This pattern contrasts with representations of the Ordovician part of the diversity trajectory shown in Sepkoski’s Phanerozoic-wide compilations. In both generic-level plots (see Sepkoski 1997: figure 1-1, 1998: figure 2) and familial-level plots (see Sepkoski 1979: figure 7, 1984: figure 1, 1997: figure 1-2), the dramatic, exponential rise of diversity is shown commencing near the beginning of the Ordovician and continuing more or less unabated—the slope of the trajectory remaining steeply inclined upward—until leveling off in the late Caradoc–early Ashgill (see Sepkoski 1984: figure 1, reproduced here as figure 1.1A).

The first major diversification pulse of Paleozoic groups commenced early in the Mid Ordovician (mid Arenig—TS.3a–b) and continued to an initial early Darriwilian (late Arenig—TS.4a) peak; then there was a short-lived mid Darriwilian (or Llanvirn—TS.4b) lag that Sepkoski linked to the North American Middle Ordovician Sauk-Tippecanoe sequence boundary (an interval, or intervals, of lowered sea level highstands; see Ross and Ross 1995; Golonka and Kiessling 2002). The second major diversification started during the late Darriwilian (formerly latest Llanvirn–early Llandeilo—TS.4c) and reached its culmination in a mid Caradoc (TS.5b–c) maximum. A limited decline followed in the late Caradoc (TS. 5d), before rapid increase resumed to the sharp, third and highest diversity peak during the early to mid Ashgill (TS.6a–b). The late Ashgill mass extinctions then caused the dramatic decline of the diversity, virtually back to levels previously attained in the early Mid Ordovician, near the start of the first of the three great diversification pulses of the Ordovician Radiation, some 25 m.y. earlier.

The second feature of Sepkoski’s (1995) documentation is the presentation of individual plots for some of the major taxonomic groups, the Cambrian EF represented by two groups (trilobites and inarticulated brachiopods), the Paleozoic EF by six groups, (articulated brachiopods, echinoderms, corals, bryozoans, nautiloid cephalopods, and rostroconch mollusks), the Modern EF by two groups (gastropods and bivalves), and two other mainly pelagic groups (graptolites and conodonts) that Sepkoski assigned to the Paleozoic EF. Some features of these patterns are outlined here, but detailed comparisons of his curves with counterparts documented in chapters of this book are not attempted because of the differing fossil data assemblies, taxonomic categories, and methods of presentation employed by many authors.

For the Early Ordovician record of more than 550 genera, the trilobite and inarticulated brachiopod components of the Cambrian EF make up about 49 percent of the total; the articulated brachiopod, nautiloid, rostroconch, bryozoan, graptolite*, and conodont* components of the Paleozoic fauna constitute about 42 percent of the total (14 percent of them pelagics—the groups with asterisks); and the gastropod and bivalve components of the Modern fauna make up about 8 percent of the total. Of these various groups, the rostroconchs have their main diversification through the Late Cambrian to Tremadocian interval, so this is one small Paleozoic group that started to diversify in the Late Cambrian, reaching its maximum in the Early Ordovician. Among other Paleozoic groups, at least one order (Ellesmerocerida) of nautiloids diversified in successive pulses across the Late Cambrian to Tremadocian boundary, and the pelagic graptolites and conodonts also radiated markedly during the Early Ordovician. This latter fact suggested to Sepkoski that, in the early history of the Paleozoic EF, some decoupling of the pelagic realm from the benthic realm may have occurred.

The marine generic diversity doubled to about 1,200 genera during the Mid Ordovician, and there were significant changes to the proportions of the three EFs. The Cambrian EF (trilobites and inarticulated brachiopods) was now represented by only about 30 percent of the total, whereas the Paleozoic EF (articulated brachiopods, echinoderms, corals, bryozoans, cephalopods, rostroconchs, graptolites*, and conodonts*) had expanded to about 62 percent of the total (nearly 12 percent of them pelagics). The Modern EF (gastropods and bivalves) remained a minor component, as in the Early Ordovician, constituting close to 8 percent of the total. A significant feature of the Paleozoic EF is that benthic elements (especially articulated brachiopods, cephalopods, echinoderms, and bryozoans) expanded more rapidly than the pelagic components through the Mid Ordovician. On the other hand, of all the groups, only the graptolites and cephalopods are shown attaining their maximum generic diversities during the Mid Ordovician. Frey et al. (chapter 21), however, show the cephalopods as reaching a relatively slightly higher maximum diversity in the Late Ordovician.

The Late Ordovician record comprises two greatest diversity increases, the first as shown in the broad, gently arching diversity curve of increase to a mid Caradoc maximum of about 1,600 genera, and the second in a higher, sharper, mid Ashgill peak of nearly 1,800 genera (figure 1.1B). It seems that all the groups surveyed by Sepkoski (1995), except apparently the rostroconchs and graptolites, attained their highest Ordovician generic diversities during this epoch. The Cambrian EF (trilobites and inarticulated brachiopods) had now declined further to about 23 percent of the total, while the Paleozoic EF (articulates, echinoderms, corals, bryozoans, cephalopods, rostroconchs, graptolites*, and conodonts*) had expanded further to about 64 percent of the total (of which nearly 8 percent were pelagics). In addition, the Modern EF (gastropods and bivalves) had now increased slightly in importance, to 12 percent of the total. All groups exhibit a sharp major decline associated with the end Ordovician extinctions.

The Late Ordovician diversity peaks of individual groups show some interesting patterns, with all the maxima associated with the early Caradoc (TS.5a), mid Caradoc (TS.5b–c), and/or the mid Ashgill (TS.6b) intervals. For the Cambrian EF groups, the trilobites have a diversity peak in the early Caradoc, while the inarticulated brachiopods have a mid Caradoc peak, though both groups exhibit a small, secondary mid Ashgill peak. Three of the Paleozoic EF groups (articulates, echinoderms, and bryozoans) show rather similar-sized mid Caradoc and mid Ashgill diversity peaks; the conodonts also show these same peaks, though they are less well defined. The nautiloids and graptolites, on the other hand, have their two prominent diversity peaks spaced farther apart, the first in the early Caradoc and the second in the mid Ashgill (TS.6b). The rostroconchs have only one minor early Caradoc peak, while the corals show only one prominent, major mid Ashgill peak. Of the components of the Modern EF, the bivalves show all three diversity peaks (early and mid Caradoc and mid Ashgill), and the gastropods show a steadily rising diversity curve to a single mid Ashgill peak.

Floral Patterns

The Ordovician floral groups have received comparatively little attention. The calcified benthic marine Ordovician algae were originally grouped by Chuvashov and Riding (1984) in their Ordovician Flora. This was differentiated as one of the three major Paleozoic evolutionary floras based on characteristic Cambrian, Ordovician, and Carboniferous assemblages associated with reefs, stromatolites, oncoids, or debris in the shallow marine carbonates. The Cambrian flora was dominated by cyanophytes (blue-green algae) that appeared during a brief span (5 m.y.) of the earliest Cambrian. The blue greens are now commonly referred to cyanobacteria because they are prokaryotic and therefore related to bacteria (Riding 1991) but differ in having chlorophyll a and in developing a typical thallus without roots, stems, or leaves (for further discussion, see chapter 31). Riding (2000) outlined the cyanophyte radiation of the Cambrian flora, from its dramatic appearance in the earliest Cambrian through its progressive decline during the Mid to Late Cambrian into the Early Ordovician, when it finally more or less disappeared.

The Ordovician flora—represented by mineralized (calcified) thallophytes—was dominated by dasyclads, codiaceans/udotaceans, and solenoporans (green and red algae). Chuvashov and Riding (1984) indicated that this flora appeared through the first two-thirds of the Ordovician Period. However, the major radiations of these groups took place later, mainly through Darriwilian to Caradoc time (see chapter 31). The great expansions of metazoans and calcified algae (Paleozoic fauna and Ordovician flora) had a profound impact on the Cambrian cyanophytes. Their decline is well documented in the reef habitat (Rowland and Shapiro 2002: figure 8; Webby 2002), but it is not yet known whether the disappearances relate to real extinctions or merely represent the loss of a preserved record when cyanophyte calcification processes ceased in unsuitable environmental conditions, possibly related to temperature decline (Riding 1992).

The other floral groups are represented by non-mineralized, microscopic thallophytes, the acritarchs, and the dispersed spores of the first bryophytelike land plants. The thallophytes are unicellular green algae (e.g., prasinophytes) and form a small component of the organic-walled phytoplankton in the Ordovician oceans. The acritarchs represent a major, organic-walled phytoplankton component of the oceans, but their identification as cysts of unicellular thallophytes remains uncertain (chapter 32). The cryptospores are predominantly dispersed spores and probably derived from small nonvascular bryophytelike plants (chapter 33). The acritarchs had an important early record in Proterozoic-Cambrian oceans—in the Early Cambrian up to 100 species (Vidal and Moczydłowska-Vidal 1997)—and then became significantly more diversified and abundant during the Ordovician (some 250 genera and about 1,300 species; table 1.1). Servais et al. (chapter 32) indicate that well-defined warmer water and cooler provincial assemblages were developed during the Early to Mid Ordovician. Based on British species data, there was apparently a progressive rise through the Early Ordovician to a Darriwilian diversity high, then continuous decline during the Late Ordovician. This contrasts with the Late Ordovician record in Baltoscandia, where Kaljo et al. (1996) have shown fluctuating, relatively high levels of diversity through the same interval, including localized peaks of diversity—two during the mid Caradoc and another in the early Ashgill. These somewhat divergent regional Mid to Late Ordovician acritarch pelagic diversity results do not closely match the record of mainly Darriwilian to Caradoc radiation events for the benthic calcified algae.

Two separate diversification phases seem to have been responsible for the earliest land-based floras. The first is indicated by the appearances of moderately abundant and cosmopolitan cryptospore records from the Darriwilian onward, and the second is suggested by the record of trilete spores in the latest Ordovician, which possibly signals the emergence of the earliest vascular plants (chapter 33). From this timing, only the first of these events can be related to the main radiation events for the benthic, calcified algae. However, it seems more likely that all these different Ordovician floral radiation events occurred completely independently of one another in their separate benthic, pelagic, and terrestrial realms.

Evolutionary Faunas at the Ecologic Level

North American and Other Continental Platforms

Sepkoski (1981b, 1991a), Sepkoski and Sheehan (1983), and Sepkoski and Miller (1985) highlighted the close relationships that exist between global diversity change within the successive Evolutionary Faunas (EFs) and localized to regional community-based diversity change. The sequential diversifications of the three EFs were recognized in the onshore-offshore patterns of expansion of the three basic trilobite-, brachiopod-, and mollusk-dominated community types. These were seen to be environmentally controlled and governed by successive onshore originations and offshore expansion, with replacement, or displacement, through time. Sepkoski and Sheehan (1983), Sepkoski and Miller (1985), and Sepkoski (1991a) employed about 500 Paleozoic (100 of them Ordovician) level-bottom communities across the North American platform for their compilations. They first converted the faunal lists to ordinal counts of generic (or species) numbers for each community and then used cluster and factor analyses to analyze the assembled data, with assembly of the results in a series of time-environmental maps. These maps depict the major changes through Ordovician time: (1) the more complexly structured Paleozoic benthic communities displacing preexisting, less-structured Cambrian communities to the outer shelf and slope; and (2) the Modern mollusk-rich communities originating onshore and, in turn, displacing the Paleozoic-dominated communities to more offshore sites.

Consequently, the Ordovician Radiation, with its significant components of the Paleozoic and Modern EFs in the brachiopod-rich and mollusk-rich communities, exerted a profound impact on the ecologic structure, with the older (Cambrian-type), and to a lesser extent the Paleozoic-type, communities being displaced diachronously offshore (Sepkoski and Sheehan 1983; Sepkoski and Miller 1985; Sepkoski 1991a). These events triggered great benthic community restructurings in the low latitudes of the North American continental shelf to slope during the Ordovician, but we are still some way from settling whether they were part of an overall global restructuring across the full range of geotectonic settings, facies profiles, and paleolatitudes or strictly regional (North American) events. Only by establishing similarly intensive, wide-ranging, and rigorous programs of Ordovician biodiversity study in other major platform regions such as Baltoscandia and South China can we necessarily expect to determine convincingly whether global processes were ultimately responsible for triggering these restructurings.

On the other hand, Miller (1997a) has shown that promising results are attained if the global patterns are dissected—that is, if diversification patterns are compared at different geographic and environmental scales. His comparative database survey of more than 6,570 genus-level Ordovician occurrences of major faunal representatives of each of Sepkoski’s (1981a) three EFs—trilobite-, brachiopod-, and mollusk-dominated—across six different continental blocks (Laurentia, Baltoscandia, North China, South China, Bohemia, and East Avalonia) revealed distinctive continent-to-continent differences in generic richness and compositional changes between the faunas through Ordovician time. Both raw data and rarefaction analyses were used, the latter to compensate for the uneven sample sizes—each bin equivalent to one very unequal British series subdivision of time. Further comparative analysis of the Laurentian and South Chinese data was provided by Miller and Mao (1998) outlining the generic diversity patterns: (1) generally higher in Laurentia than in South China; (2) levels rather static in the Early to Mid Ordovician, then becoming higher in the Caradoc for Laurentia; (3) levels in the Mid and Late Ordovician probably somewhat higher than for the Early Ordovician of South China; and (4) increases of benthic mollusks only in the Late Ordovician of Laurentia. The major contrast in the patterns of diversification of the two continental areas relates to the presence or absence of siliciclastic-rich environments. Carbonate sedimentation was predominant in South China, but there were influxes of terrigenous sediments across eastern and central Laurentia during the Late Ordovician, associated with orogenic activity (Taconic Orogeny), and this facilitated the radiation of benthic mollusks, especially the bivalves (chapter 20). Miller and Mao (1998) recognized that