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Y Y A A P P OFFICERS AND EDITORS OF THE PALEONTOLOGICAL SOCIETY Grand Patrons ANDREW H. KNOLL RICHARD K. BAMBACH CHARLES LANE ARTHUR J. BOUCOT ALLAN R. LARSON President PETER J. HARRIES, SIMON SCHNEIDER, Editor, Special Publications RODNEY M. FELDMAN AND CARRIE SCHWEITZER CECILIA LENK AND PAUL K. STROTHER BRUCE MacFADDEN, North Carolina State CASP COLIN SUMRALL, ROBERT AND ELVIRA GASTALDO JERE H. LIPPS Florida Museum of Natural History CRAIG SCOTT, Univ. of Tennessee, Knoxville THE FAMILY OF N. GARY LANE ALAN R. LORD MICHAEL HAUTMANN, bmacfadd@fl mnh.ufl .edu Royal Tyrrell Museum of Paleontology [email protected] A. R. (PETE) PALMER CHRIS MAPLES AND SARA MARCUS Univ. of Zurich J. WILLIAM SCHOPF President-Elect MARY SILCOX, Education/Outreach Coordinator JANET WOOD McGREGOR RAMAN J. SINGH WILLIAM DiMICHELE, MARTIN J. HEAD, Univ. of Toronto Scarborough ROWAN LOCKWOOD, CHERYL L. METZ STEVEN M. STANLEY Smithsonian Institution Brock Univ. College of William & Mary MOLLY F. AND CALVIN MILLER EDUARDO LEORRI SORIANO, [email protected] [email protected] RONALD W. MORIN THOMAS HEGNA, East Carolina Univ., Greenville Patrons OSBORNE B. NYE, JR. Past President Western Illinois Univ. Diversity and Inclusion Coordinator EDWARD J. ANDREW, JR. DAVID VARRICCHIO, TERESA O’NEILL ARNOLD I. MILLER, DENA SMITH, WILLIAM I. AUSICH Montana State Univ. RONALD L. PARSLEY Univ. of Cincinnati SAMANTHA HOPKINS, Univ. of Colorado BRUCE M. BELL MARK E. PATZKOWSKY [email protected] Univ. of Oregon STEVE WESTROP, [email protected] STIG M. BERGSTROM JOHN POJETA, JR. Oklahoma Museum DAVID J. BOTTJER MARY LOU POJETA Secretary NATHAN JUD, Amateur/Avocational Liaison DONALD W. BOYD JOHN K. POPE CLAUDIA JOHNSON, Univ. of Florida DAVID M. WORK, JAYSON KOWINSKY, DEREK E. G. BRIGGS ANNE RAYMOND Indiana Univ. Maine State Museum Fossilguy.com JAMES C. BROWER DANIEL KSEPKA, MARGARET N. REES [email protected] [email protected] JOHN L. CARTER Bruce Museum FRANK H. T. RHODES PATRICK WYSE JACKSON, PETER R. CRANE RICHARD A. ROBISON Treasurer Univ. of Dublin Trinity College Representatives At Large THE CUSHMAN FOUNDATION JAMES LAMSDELL, JUNE R. P. AND CHARLES A. ROSS ALYCIA STIGALL, GABRIELA MÁNGANO, J. ROBERT DODD West Virginia Univ. MARGARET M. YACOBUCCI, CHRISTOPHER A. SHAW Ohio Univ. Univ. Saskatchewan MARY L. DROSER [email protected] Bowling Green State Univ. [email protected] FREDERICK C. SHAW MAX LANGER, LUCY E. EDWARDS SAMUEL ZAMORA, LAWRENCE M. SMALL JP Editors Universidade de São Paulo CAROLINE STRÖMBERG, ROBERT J. ELIAS Geological Survey of Spain GEORGE D. STANLEY, JR. BRENDA R. HUNDA, Univ. of Washington J. MARK ERICKSON BERTRAND LEFEBVRE, DOUGLAS H. ERWIN ROBERT J. STANTON, JR. Cincinnati Museum Center RENBIN ZHAN, [email protected] Lyon University HOWARD R. FELDMAN CARL W. STOCK [email protected] Nanjing Institute of Geology and ROBERT M. FINKS ROGER D. K. THOMAS Palaeontology, Student Representative STEPHEN A. LESLIE, KARL W. FLESSA BRUCE H. TIFFNEY JISUO JIN, Chinese Academy of Sciences KRISTOPHER M. KUSNERIK, Univ. of Western Ontario James Madison Univ. Univ. of Florida TERRENCE J. FREST SALLY E. WALKER [email protected] SHUNXIN ZHANG, kmkusnerik@ufl .edu PHILIP D. GINGERICH RON F. WASZCZAK BRUCE LIEBERMAN, JOHNNY A. WATERS Univ. of Kansas Canada-Nunavut Geoscience Offi ce BRIAN F. GLENISTER HANS-DIETER SUES, BRIDGET KELLY, F. D. (BUD) HOLLAND, JR. GARY D. WEBSTER Smithsonian Institution ZHIFEI ZHANG, ALEX LIU, Univ. of California, Riverside J. STEWART HOLLINGSWORTH MARK A. WILSON [email protected] Northwest Univ. Univ. of Cambridge [email protected] NIGEL C. HUGHES SCOTT L. WING THOMAS W. KAMMER ABBY YOCHELSON ELLEN CURRANO, Paleobiology Senior Editor Section Chairs STEVEN LoDUCA, ERLE G. KAUFFMAN CHARLES YOCHELSON University of Wyoming MARK PATZKOWSKY, Cordilleran: Eastern Michigan Univ. PATRICIA H. KELLEY JEFFERY B. YOCHELSON [email protected] Penn State Univ. CARLIE PIETSCH, SUSAN M. KIDWELL LOUIS G. ZACHOS RICK LUPIA, San Jose State Univ. JP Associate Editors Paleobiology Co-Editors [email protected] JOSEPH BOTTING, Univ. of Oklahoma NADIA FRÖBISCH, Nanjing Institute of Geology GABRIELA MÁNGANO, Museum für Naturkunde North Central: Th e Journal of Paleontology publishes manuscripts on all aspects of paleontology, including systematics, phylogeny, paleoecology, paleobiogeography, biostratigraphy, and Paleontology, Univ. Saskatchewan CARRIE SCHWEITZER, taphonomy, and evolution. It emphasizes specimen-based research and features illustrations of the highest quality possible. Manuscript treating all fossil organism groups Chinese Academy of Sciences KEVIN BOYCE, Kent State Univ. Stanford Univ. are welcome, including invertebrates, vertebrates, plants, algae, and microorganisms of all sorts, as well as ichnofossils (trace fossils). GREG McDONALD, [email protected] STYLIANOS CHATZIMANOLIS, Bureau of Land Management WOLFGANG KIESSLING, Univ. Tennessee–Chattanooga Northeastern: Th e Journal of Paleontology accepts only online manuscript submissions through the website: Universitat Erlangen JUDITHNA GEL-MYERS, TOM CRONIN, MALGORZATA https://mc.manuscriptcentral.com/paleo JAMES CRAMPTON, St. Lawrence Univ. U.S. Geological Survey MOCZYDLOW SKA-VIDAL, Uppsala Univ. GNA Science [email protected] ______WILLIAM A. DiMICHELE, Managing Editor Rocky Mountain: Author guidelines can be downloaded at: Smithsonian Institution RICHARD MOOI, JESSICA KASTIGAR, BROOKE LONG, California Academy of Sciences https://www.cambridge.org/core/journals/journal-of-paleontology/information/instructions-contributors JAN OVE EBBESTAD [email protected] S. Dakota School of Mines & Tech. Museum of Evolution, ALISON MURRAY, [email protected] [email protected] ______Uppsala University Univ. of Alberta Communications Offi cer South Central: For questions regarding online submissions, please contact the editorial offi ces at: ROBERT J. ELIAS, PAULA NOBLE, TONY MARTIN, VICTORIA CHRAIBI, [email protected] Univ. of Manitoba Univ. of Nevada Emory Univ. Tarleton State Univ. [email protected] [email protected] ______NADIA FRÖBISCH, ANDRZEJ PISERA, Southeastern: Museum für Naturkunde Polish Academy of Sciences Program Coordinator Membership Information MATTHEW CLAPHAM, MICHELLE CASEY, ELIZABETH HARPER, BRIAN PRATT, Univ. of California, Santa Cruz Murray State Univ. Members of the Paleontological Society automatically receive electronic subscriptions to Paleobiology and Univ. of Cambridge Univ. of Saskatchewan [email protected] [email protected] the Journal of Paleontology as a benefi t of membership. To apply for individual membership, please go to the Paleontological Society’s homepage at http://paleosoc.org/ where links to an electronic application, a printable paper application form, and a complete list of membership benefi ts are provided under the Membership heading. ______Cover illustration: Cyphaspides from the Bou Tchrafi ne Formation, Eifelian Age, Middle Devonian, northwest of Jorf, southeastern Morocco. Chatterton et al., Fig. 5.2. Visit Th e Paleontological Society website at www.paleosoc.org.

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