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A Revision of the Genus Strategus (Coleoptera: Scarabaeidae)

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A Revision of the Genus Strategus (Coleoptera: Scarabaeidae) University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Bulletin of the University of Nebraska State Museum Museum, University of Nebraska State 1976 A Revision of the Genus Strategus (Coleoptera: Scarabaeidae) Brett C. Ratcliffe University of Nebraska-Lincoln, [email protected] Follow this and additional works at: https://digitalcommons.unl.edu/museumbulletin Part of the Entomology Commons, Geology Commons, Geomorphology Commons, Other Ecology and Evolutionary Biology Commons, Paleobiology Commons, Paleontology Commons, and the Sedimentology Commons Ratcliffe, Brett C., "A Revision of the Genus Strategus (Coleoptera: Scarabaeidae)" (1976). Bulletin of the University of Nebraska State Museum. 41. https://digitalcommons.unl.edu/museumbulletin/41 This Article is brought to you for free and open access by the Museum, University of Nebraska State at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Bulletin of the University of Nebraska State Museum by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. BULLETIN OF VOLUME 10, NUMBER 3 The University of Nebraska State Museum FEBRUARY, 1976 Brett C. Ratcliffe <~ Revision of the Genus Strategus (Coleoptera: Scarabaeidae) Brett C. Ratcliffe A Revision of the Genus Strategus (Coleoptera: Scarabaeidae) BULLETIN OF The University of Nebraska State Museum VOLUME 10, NUMBER 9 FEBRUARY, 1976 Frontispiece.-Strategus a/oeus (L.). type of the genus. BULLETIN OF VOLUME 10, NUMBER 3 THE UNIVERSITY OF NEBRASKA STATE MUSEUM February, 1976 Pp. 93-204, Tables 1-7 Frontispiece, Figs. 1-176 ABSTRACT A Revision of the Genus Strategus (Coleoptera: Scarabaeidae) Brett C. Ratcliffe The New World genus Strategus Kirby is here defined and revised for the first time. Eight species are described as new: S. at/anticus, S. caymani, S. craigi, S. hipposiderus, S. howdeni, S. /on­ gichomperus, S. symphenax, and S. tarquinius; 12 new junior synonyms are established (includ­ ing the rejection of six previously valid species)'; one species is raised from synonymy; and two new replacement names are proposed. Strategus now contains one fossil species and 31 valid extant species. A key to the males and females of all the species is provided for the first time. All taxa are described or redescribed and illustrated by habitus figures and drawings of the male genitalia. Biologies are discussed when data are available. Extensive distributional data and locality record dot maps are presented as well as a zoogeographical analysis of the genus. Lastly, a computer-assisted cladistic reconstruction of the presumed phylogeny of the genus is provided. CONTRIBUTION OF The Division of Entomology of the University of Nebraska State M\Jseum and Contribution Number 395, Department of Entomology, University of Nebraska. Copyright © 1976 by the Board of Regents of the University of Nebraska Library of Congress Catalog Card Number 76-1109 ISSN 0093-6812 Manufactured in the United States of America Ratcliffe1 A Revision of the Genus Strategus (Coleoptera: Scarabaeidae) ment of the genus to date, but did not include a ..fTRODUCTION key or describe all the species known to him. The genus Strategus was established by Wil­ In 1915 Casey erected Anastrategus as a new liam Kirby in 1828 when he listed those species genus in the Pentodontini to include those to be included in the new genus; these were S. species of former Strategus in which the males a/oeus (L.), s. antaeus (Drury), S. syphax (Fabr.), lacked horns, i. e., S. adolescens Kolbe, S. ces­ and S. titan us (Fabr.) ( = S. simson (L.)). A sub­ sus LeConte (and those synonyms of Casey and sequent definition of what characterized the Kolbe), S. fallaciosus Kolbe, and S. splendens genus based on all the included species has (Beauvois) (and those synonyms of Casey). never been given although Burmeister (1847) Arrow (1937a) synonymized the genus with made a good attempt and Paulian (1947), Strategus and Endr6di (1959) resurrected Ana­ LeConte (1861-62), and Saylor (1946) estab­ strategus as a subgenus of Strategus and in­ lished partial descriptions. The genus as a whole cluded in it the same species as had Casey. has not been treated in a monograph nor has Casey (1915) also proposed Strategodes as a there ever been a dichotomus key to all of the subgenus of Strategus to incorporate those species. Most of those species descriptions species that lack a sutural stria and have long, given in the past have been very short and vague. slender mandibular teeth; these included S. an­ Regional works with keys to the local species taeus (Drury) (and synonyms of Casey) and S. (primarily the U. S. species) have been provided mormon Burmeister. Chapin (1932a) established by Casey (1915), Chapin (1932a and b), Dillon Strategopsis as a subgenus for those species and Dillon (1961), Horn (1875), Paulian (1947), with an unarmed or unidentate galea; this in­ and Saylor (1946). Checklists for Strategus have cluded S. sarpedon (Burmeister). been given by Arrow (1937a), Blackwelder Subgenera are not recognized in this revision (1944), Bruch (1911), Dejean (1836), Fleutiaux as I do not believe there exist valid reasons for and Salle (1889), Gemminger and Harold (1869), doing so. Certainly Casey's Anastrategus, based Henshaw (1885), Kolbe (1906), Leng and Mutch­ on a variable male sexual character, cannot be ler (1914, 1917), Leng (1920), and Wolcott (1923, justified as Arrow (1937b) has previously pOinted 1936). Synonymies were somewhat stabilized by out. Casey's Strategodes and Chapin's Arrow (1937a) and the last species described as Strategopsis are each based on a single derived new was also established by Arrow (1947). Kolbe and ancestral character state, respectively, and (1906) has given the most comprehensive treat- do not reflect probable phyletic relationships. Only moderate and sketchy biological data are 'Systematics Collections, W-436 Nebraska Hall, University actually known for Strategus species even of Nebraska State Museum, Lincoln, Nebraska 68588. though the majority probably have similar life 94 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM cycles. Distributional information for the species California Academy of Sciences (CASC); Hugh B. has been noticeably fragmentary and erroneous, Leech. and, even after the results of this study, could Canadian National Collection of Insects (CNCI); still be considerably refined. J. M. Campbell, Henry F. Howden. Strategus has been characterized by Endrodi Carnegie Museum of Natural History (ICCM); (personal communication) as the most taxonom­ George E. Wallace. ically difficult genus of Oryctini in the New World Field Museum of Natural History (FMNH); Henry next to Heterogomphus because of its high level S. Dybas, Michael Prokop. of intraspecific variation. This state of affairs has Illinois Natural History Survey (INHS); Milton W. undoubtedly discouraged any comprehensive Sanderson. systematic treatment in the past. While concur­ Kansas State University (KSUC); H. Derrick ring with Dr. Endrodi's analysis, it is hoped that Blocker. this study will sufficiently and accurately inter­ Los Angeles County Museum of Natural History pret the genus so that it will no longer be the (LACM); Roy R. Snelling. source of confusion that it has been. Museo de Historia Natural de la Ciudad de This systematic treatment delineates the Mexico (MHNCM); Alfredo Barrera. genus and provides keys to the males and Museum National d'Histoire Naturelle, Paris .females of all the species for the first time. It (MNHN); A. Descarpentries. establishes complete synonymies, provides ex­ tensive distributional data, describes each Museum of Comparative Zoology (MCZC); Frank species, brings together most, if not all, the pub­ M. Carpenter, Charles G. Oliver, Janice C. Scott. lished biological information for the species, and presents a zoogeographic analysis of the genus. National Museum of Natural History (USNM); All species and all distributions are illustrated. Robert D. Gordon. Eight species are described as new, 12 new Ohio State University (OSUC); Charles A. Triple­ synonymies are established including the rejec­ horn. tion of six previously valid species, one species Oregon State University (OSUO); PaulO. Ritcher. is raised from synonymy, and two new replace­ Peabody Museum of Natural History (PMNH); ment names are proposed. Strategus now con­ Kirby W. Brown, Charles L. Remington. tains one fossil species and 31 valid extant Santa Barbara Museum of Natural History species. (SBMN); Nelson W. Baker. Texas A & M University (TAMU); Horace R. Burke. Universidad de Concepcion, Chile (UC); Tomas Cekalovic K. ACKNOWLEDGMENTS Universidad de la Republica, Montevideo (URU); Carlos S. Carbonell. This study could not have been completed Universidad Nacional de Colombia (UNC); Rafael without the generous assistance of many indi­ Cancelado. viduals. The following institutions and staff are Universidade de Sao Paulo (USP); Ubirajara R. gratefully acknowledged for the loan of speci­ Martins. mens or assistance in studying specimens in Universitetets Zoologiske Museum, Copenhagen their care (standard abbreviations for North (UZM); N. Mfi'ller Andersen, S. G. Larsson. American collections as in Arnett and Samuel­ University of Colorado (UCMC); U. N. Lanham. son, 1969): University of Kansas (SEMC); George W. Byers. University of Michigan (UMMC); Thomas E. Moore. American Museum of Natural History (AMNH); University of Nebraska (DEUN). Lee H. Herman, Jr. University of Wisconsin (UWEM); John R. Baker. British
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