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Bulletin of the Peabody Museum of Natural History Bulletin of the Peabody Museum of Natural History In publication since 1925, and originally a monograph series, the Bulletin of the Peabody Museum of Natural History publishes peer-reviewed contributions on original research in the natural sciences represented by the collections of the Yale Peabody Museum of Natural History’s curatorial divisions, covering diverse topics that include evolution, phylogeny, taxonomy, systematics, biology, botany, zoology, invertebrate and vertebrate paleontology, and paleoecology, paleobotany, and archaeology. Full monographs of Bulletin numbers 1 through 46 are available for download at peabody.yale.edu. Beginning with Volume 47, fully indexed published Bulletin articles are available online through BioOne Complete. Yale University provides access to these materials for educational and research purposes only. Copyright or other proprietary rights to content contained in this document may be held by individuals or entities other than, or in addition to, Yale University. You are solely responsible for determining the ownership of the copyright, and for obtaining permission for your intended use. Yale University makes no warranty that your distribution, reproduction, or other use of these materials will not infringe the rights of third parties. This work is licensed under the Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-sa/4.0/ or send a letter to Creative Commons, PO Box 1866, Mountain View, CA 94042, USA. PEABODY MUSEUM OF NATURAL HISTORY, YALE UNIVERSITY 170 WHITNEY AVENUE, P.O. BOX 208118, NEW HAVEN CT 06520-8118 USA PEABODY.YALE.EDU Systematic Revision of the Olenelloidea (Trilobita, Cambrian) Olenellus getzi Dunbar [YPM 6657], holotype. Early Cambrian, Pennsylvania, U.S.A. Copyright © 1997 Peabody Museum of Natural History, Yale University. Photograph by William K. Sacco. Bulletin of the Peabody Museum of 45Natural History Yale University 18 October 1999 New Haven, Connecticut Systematic Revision of the Olenelloidea (Trilobita, Cambrian) Bruce S. Lieberman, Ph.D. Department of Geology University of Kansas Curatorial Affiliate Division of Invertebrate Paleontology Peabody Museum of Natural History Yale University BULLETIN OF THE PEABODY MUSEUM OF NATURAL HISTORY Curatorial Editor-in-Chief Jacques A. Gauthier Executive Editor Lawrence F. Gall Publications Editor Rosemary Volpe The Bulletin of the Peabody Museum of Natural History, Yale University, publishes independent monographs in the natural sciences based on research in the collections of the Yale Peabody Museum. Volumes are numbered consecutively and appear at irregular intervals. The Bulletin incorporates the Bulletin of the Bingham Oceanographic Collections, which ceased independent publication after Volume 19, Article 2 (1967). The series is now out of print. For submission guidelines and information on ordering Yale Peabody Museum publications, see our website or contact: Publications Office Peabody Museum of Natural History Yale University 170 Whitney Avenue P.O. Box 208118 New Haven, CT 06520-8118 U.S.A. Phone: (203) 432-3786 Fax: (203) 432-9816 E-mail: [email protected] URL: http://www.peabody.yale.edu/ On the cover: Homotelus bromidensis [YPM 10186]. Early Ordovician, Oklahoma, U.S.A. Copyright © 1997 Peabody Museum of Natural History, Yale University. Photograph by B.S. Lieberman. Copyright © 1999 Peabody Museum of Natural History, Yale University. All rights reserved. This book may not be reproduced, in whole or in part, including illustrations, in any form (except by reviewers for the public press) without the written permission of the publisher. ISBN: 0-912532-47-5 Printed in the U.S.A. ∞ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). CONTENTS vii List of Figures viii List of Tables ix Abstract ix Keywords x Acknowledgments 3 Introduction 7 one • Phylogenetic Analysis of the Olenellinae 8 Systematic Paleontology 13 Included Taxa 13 Olenellus 27 two • Phylogenetic Analysis of the Mesonacinae 28 Systematic Paleontology 32 Included Taxa 32 Mesolenellus 38 Mesonacis 47 three • Phylogenetic Analysis of Elliptocephala 49 Systematic Paleontology 49 Included Taxa 49 Wanneria 53 Elliptocephala 65 four • Phylogenetic Analysis of the Holmiidae 67 Systematic Paleontology 73 Included Taxa 73 Holmia 76 Baltobergstroemia new genus 80 Kjerulfia 81 Schmidtiellus 83 Palmettaspis 85 Esmeraldina 86 Holmiella 89 five • Phylogenetic Analysis of Fritzolenellus 89 Systematic Paleontology 90 Included Taxa 90 Fritzolenellus 99 six • Phylogenetic Analysis of Mummaspis 99 Systematic Paleontology 99 Included Taxa 99 Mummaspis vi Peabody Museum Bulletin 45 107 seven • Phylogenetic Analysis of the Bristoliinae 114 Systematic Paleontology 114 Included Taxa 114 Laudonia 116 Lochmanolenellus 118 Fremontella 120 Bristolia 129 eight • Phylogenetic Analysis of Bolbolenellus 129 Systematic Paleontology 131 Included Taxa 131 Nephrolenellus 134 Bolbolenellus 141 nine • Integrating Phylogenetic and Stratigraphic Data 145 References Systematic Revision of the Olenelloidea (Trilobita, Cambrian) vii FIGURES 12 Figure 1. A strict consensus of 40 most parsimonious trees of length 36 steps produced from analysis of character data in Table 2. 14 Figure 2. Olenellus spp. 18 Figure 3. Olenellus spp. 20 Figure 4. Olenellus robsonensis (Burling). 22 Figure 5. Olenellus getzi Dunbar. 24 Figure 6. Olenellus getzi Dunbar. 29 Figure 7. The most parsimonious tree of length 60 steps produced from analysis of character data in Table 4. 40 Figure 8. Mesonacis fremonti (Walcott). 44 Figure 9. Mesonacis spp. 48 Figure 10. A strict consensus of two most parsimonious trees of length 53 steps produced from analysis of character data in Table 6. 54 Figure 11. Wanneria walcottana (Wanner); Holmia kjerulfi (Linnarsson). 56 Figure 12. Elliptocephala spp. 66 Figure 13. A strict consensus of the six most parsimonious trees of length 90 steps produced from analysis of character data in Table 8. 68 Figure 14. The weighted tree of length 32,490 steps, with the same topology as one of the most parsimonious trees used to derive the consensus tree shown in Figure 13. 79 Figure 15. Esmeraldina rowei (Walcott); Palmettaspis consorta Fritz; Holmiella falcuta Fritz. 90 Figure 16. The most parsimonious cladogram of length 5 steps produced from analysis of character data in Table 10. 94 Figure 17. Fritzolenellus truemani (Walcott); Mummaspis occidens (Walcott); Mummaspis truncatooculatus (Fritz); Mummaspis muralensis (Fritz); Mummaspis macer (Walcott). 100 Figure 18. The most parsimonious cladogram of length 9 steps produced from analysis of character data in Table 12. 108 Figure 19. A strict consensus of the four most parsimonious cladograms of length 35 steps produced from analysis of character data in Table 14. 112 Figure 20. Laudonia bispinata Harrington; Bristolia insolens (Resser); Bristolia harringtoni, new species; Bristolia mohavensis (Hazzard and Crickmay); Nephrolenellus jasperensis, new species; Bolbolenellus groenlandicus (Poulsen). 130 Figure 21. The most parsimonious tree of length 14 steps produced from analysis of character data in Table 19. 137 Figure 22. Bolbolenellus groenlandicus (Poulsen); Bolbolenellus altifrontatus (Fritz). 142 Figure 23. A phylogenetic tree depicting topology and timing of patterns of speciation in the Olenelloidea. viii Peabody Museum Bulletin 45 TABLES 10 Table 1. Description of characters and character states used in phylogenetic analysis of Olenellus. 16 Table 2. Character state distributions for Olenellus and outgroup used in phylogenetic parsimony analysis. 30 Table 3. Description of characters and character states used in phylogenetic analysis of Mesonacinae. 34 Table 4. Character state distributions for Mesonacinae and outgroups used in phylogenetic parsimony analysis. 50 Table 5. Description of characters and character states used in phylogenetic analysis of Elliptocephala. 52 Table 6. Character state distributions for Elliptocephala and outgroup used in phylogenetic analysis. 70 Table 7. Description of characters and character states used in phylogenetic analysis of Holmiinae. 76 Table 8. Character state distributions for Holmiinae and outgroup used in phylogenetic analysis. 92 Table 9. Description of characters and character states used in phylogenetic analysis of Fritzolenellus. 93 Table 10. Character state distributions for Fritzolenellus and outgroups used in phylogenetic analysis. 102 Table 11. Description of characters and character states used in phylogenetic analysis of Mummaspis. 104 Table 12. Character state distributions for Mummaspis and outgroup used in phylogenetic parsimony analysis. 110 Table 13. Description of characters and character states used in phylogenetic analysis of Bristoliinae. 115 Table 14. Character state distributions for Bristoliinae and outgroups used in phylogenetic parsimony analysis. 122 Table 15. The first two principal components generated from analysis of morphometric data from species of the genus Bristolia. 124 Table 16. A summary of classification produced from a linear discriminant analysis. 126 Table 17. The results of a linear discriminant analysis. 133 Table 18. Description of characters and character states used in phylogenetic analysis of Bolbolenellus. 135 Table 19. Character state distributions for Bolbolenellus and outgroups used in phylogenetic parsimony analysis. Systematic Revision of the Olenelloidea (Trilobita, Cambrian) ix ABSTRACT
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