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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]

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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 Museum (Natural History) (BMNH); M. E. Zoologisches Museum der Humboldt- Bacchus, R. D. Pope. Universitiit, Berlin (ZMHU); F. Hieke. A REVISION OF THE GENUS Strategus I 95

· ens were also seen from the following support, and guidance of Kenneth P. Pruess Speclm .. (Department of Entomology, University of Ne­ private collections. braska) under whose supervision this study was John D. Glaser (JOG), Baltimore, Maryl~nd .. conducted. Alan R. Hardy (AHCC), Sacramento, California. Henry F. Howden (HAHC), Canada. ~ttawa, METHODS Brett C. Ratcliffe (BCRC), Lincoln, Nebraska, Richard L. Westcott (RLWE), Salem, Oregon. The results of this study were based on the examination of 5,115 specimens from 29 institu­ Deep appreciation is also ex~ended to: AAnto~io tional and five private collections (see Acknow­ eeltr~n R. (Instituto Colom~l~no Agropecu~lo, ledgments). Most of the major collections in the Bogota) for providing addltlo~al. Colo":,blan United States, Canada, and Mexico were visited, _,imens a~d data;, Sebo Endrodl (Termesz~t­ and collecting trips were conducted to Kansas, ttJdomany Muzeum Allattara, Budapest) for In- New Mexico, Arizona, British Honduras (Belize), 1otmation regarding Burmeister's types; Ake Mexico, Panama, and to the common borders of Holm and Thure Palm (Uppsala Universitet, Upp­ Colombia, Brazil, and Peru. Collecting "Ia) for dissecting out and providing illustra­ techniques consisted of random searching in tl~s of the genitalia of the type of Strategus favorable areas, blacklighting, and frequenting • 'oBuS (L.); M. E. Bacchus (British Museum) for lights at night in selected locales . providing drawings and photographs of the type Early in the study an attempt was made to ob­ of Strate gus syphax (Fabr.); C. Besuchet (Musee tain Strategus larvae from numerous sources so d'Histoire Naturelle de Geneve, Geneva) for pro­ that a key could be provided for many of the im­ viding information on the type of Strategus ajax matures as well as for the adults. Even though (Olivier); F. Chalumeau and L. Gruner (Station de the larvae of some species are well known due to Recherches de Zoologie et Lutte Biologique, their past economic significance, little success Guadeloupe) for providing specimens and addi­ was had in trying to obtain larvae on loan and, tional distributional data from Guadeloupe; therefore, plans to key and describe the larvae Janet E. Howe (University of Chicago) for per­ had to be abandoned (at least for the present). mission to use the Goode Base Maps in Figs. The apparent lack of immature specimens in col­ 4-9; Martha Haack (University of Nebraska State lections was a major disappointment to the au­ Museum) for illustrating the frontispiece and thor. The reader is referred to Ritcher (1944, Figs. 174-175. 1966) for keys and descriptions to the larvae of Special thanks~re extended to Henry F. How­ Strategus aloeus (L.) ( = julian us Burmeister), S. den (Carleton University, Ottawa) for providing antaeus (Drury), and S. splendens (Beauvois). me with helpful discussions and with the oppor­ A conventional artificial key to the males and tunity to collect with him in South America and females of all the species is presented for the to Charles D. Lofton (McCook, Nebraska) for his first time. Although many morphological charac­ faithful collecting assistance during trips to Cen­ ters within the genus are highly labile, an attempt tral and South America. was made to utilize key characters which were This study was supported in part by travel consistantly expressed, low in intrinsic variabil­ funds from the University of Nebraska Research ity, and easily observed with reasonable proce­ Council, Society of the Sigma Xi, Myron H. dures. Swenk Memorial Fund (Department of Entomol­ The morphological characters used in this ogy, University of Nebraska), the University of study were selected because they were seen to Nebraska State Museum, Department of Ento­ vary on a consistent basis between phena, thus mology (University of Nebraska), and a Warren F. allowing for separation of different taxa based and Edith R. Day Dissertation Travel Fellowship on the character state. Phena were sorted ac­ (University of Nebraska). cording to the following characters: length and .L.a~tly, I am deeply grateful for the constructive width measurements; sculpturing and hairiness Criticism, encouragement, numerous letters of of the front; shape and sculpturing of the 96 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

clypeus; shape and size of the mandibular lobes; scriptions are g:ven for the males and females of size of the head tubercles; interocular width; each species for the first time, and all observed sculpturing of the mesosternum and pronotum; variation in the character states is recorded. form and size of the horns; sculpturing of the Technical terminology is essentially that of elytra; the shape, hairiness, and sculpturing of Torre-Bueno (1937). For greater convenience the the pygidium; and the form of the parameres of species descriptions and illustrations are ar­ the male genitalia. Sorting was not done on a ranged alphabetically. geographical basis except in the rare instances Biologies or life history notes, when known, of morphologically inseparable females. are included in the discussion of the appropriate Taxa were initially defined a priori within the species. limits of existing species descriptions. The use of Locality records are presented in the following additional characters never before used and the format: country; state, province, or equivalent; re-evaluation of previously recognized charac­ locale. A temporal distribution follows the rec­ ters in much larger samples sizes and in the ords for each country. A few specific literature types enabled me to redefine or more precisely references and personal communications citing limit the definition of the various species. New distributional data are used sparingly and are species were recognized when character states appropriately acknowledged, but checklists were seen to fall significantly outside the range were not used. Location and correct spellings of of my more comprehensive redefinitions of exist­ geographic place names were verified by con­ ing taxa. sulting American Geographical Society (1943a­ Several characters were disregarded in de­ c, 1944a-i) and Bartholomew (1957). Distribu­ lineating taxa because they were either constant, tional dot maps are provided for each species too variable, or examples of random character (except S. adolescens Kolbe, for which no states. These were: shape and sculpturing of the specific locales are known). mentum (two exceptions), palps, antennae, and Dorsal and lateral habitus figures of the male eye canthus; punctation of the mandibles; gen­ of each species are presented by means of eral punctation of the horns (two exceptions); photographs; the females are not similarly illus­ curvature of the lateral margin of the pronotum; trated because no appreciable differences would shape and sculpturing of the scutellum; angle of be noticed among them. Lateral and caudal the elytral apices; size of the stridulatory ridges views of the male genitalia of each species are on the propygidium; punctation and hairiness of illustrated by line drawings made with the use of the abdominal sternites; shape of the female camera lucida. Specimens were observed with a genital plates; form and sculpturing of the apices Leitz stereomicroscope using magnifications of of the tibiae; length and ratios of tarsal seg­ 12.5 and 50.0 x. ments; and size and shape of claws. Complete synonymies are provided for all the EXPLANATION OF CHARACTERS species. The historical usage of names within USED IN KEYS AND DESCRIPTIONS each species is not given in the synonymy sec­ tion as is frequently customary. I believe such Measurements: Length measurements were usage may tend to obfuscate actual synonymies; made from the apex of the clypeus to the apex if different usages of names are significant, they of the elytra and width measurements were are mentioned in the text. It should be noted that made across the humerus. Fabricius often knowingly redescribed in later Clypeus: Several authors (notably Bates, 1886- works species that had been previously named. 1890; Burmeister, 1847; Kolbe, 1906) have re­ Those redescriptions listed in the synonymies in ferred to the clypeus and its apical shape as this paper appeared without references to any this is of considerable importance for correct prior publications and could technically, there­ identifications. From their descriptions, it is fore, have been taken as new species descrip­ apparent that these authors were, in most in­ tions although, of course, they were not. stances, referring to the form of the apex of the Reasonably complete and comparable de- clypeal disc and not to the entire clypeus. If • A REVISION OF THE GENUS Strategus / 97

only the apex of the disc is considered, it is als within a species with well-developed arma­ possible to have an apical shape considerably ture which is possibly the result of an optimum different from that of the entire clypeus which nutritional history in the larval stage. Minors also includes the declivous sides. The apex of are those individuals within a species whose the clypeus, then, is the true apex and not the armature has not developed to the recognized apex of the clypeal disc alone. Care should potential development of that particular also be taken when arriving at a conclusion species. There is no clear boundary separating about the form of the clypeal apex as this area the two forms of development and all grada­ is very subject to having its shape altered be­ tions between the two extremes occur. cause of abrasion resulting from digging by Elytra: In those species with rows of punctures the beetle. Fortunately, a worn clypeus can on the disc, the rows are numbered from the usually be recognized as such and so not be sutural stria laterally to the humerus. The rows misinterpreted. on the sides of the elytra are numbered from Mentum: At first thought to be a good character, the median (humeral) border of the sides later­ the mentum was subsequently found to be in­ ally to the elytral margin. consistently variable in regard to shape and Pygidium: The basal, transverse band of setiger­ sculpturing of the disc and shape of the apex. ous punctures or rugosity is of little taxonomic With rare exception, this character is not used. value and is not considered in the species de­ Mandibles: The form of the three lobes of each scriptions. The setae referred to on the apical mandible is of taxonomic significance. margins mean those setae inside the marginal Whereas the first and third lobes do not usu­ bead of the pygidium and not those just poste­ ally vary to any great extent, the middle lobe is rior of the marginal bead. The lateral emargi­ differentially developed among many species nation is the oblong, shallow, excavated, or and is often diagnostic. The mandibles are depressed area near the margins on either side subject to considerable wearing and, as with of the apex; it is frequently absent or nearly the clypeal apex, judicious care should be em­ obsolete. ployed when ascertaining the shape of the Genitalia: The shape of the parameres of the lobes. male genitalia is an important and rather con­ Interocular width: Measured by the number of sistant character. Some species have very transverse eye diameters necessary to span similar genitalia, and so this character should the interocular gap, this character is consis­ be used in combination with other characters tently expressed within the limits of the de­ for an accurate determination. The female scriptions. genitalia were not used in this study as a pre­ Mesosternum: nie anterior, oblique half, or the liminary examination of them revealed few dif­ entire mesosternum is setigerously punctate. ferences in the genital plates. The punctures are usually large and ocellate. Punctures: Punctures are considered irregular in Pronotum: For convenience, the descriptions di­ distribution and simple unless otherwise vide the pronotum into disc, sides, and an­ noted. Ocellate punctures are ringed with a terior third or half (which includes the fovea). slightly different color tone, and umbilicate The base of the pronotum of most species has punctures are navel-shaped or have a convex a transverse band of coarse sculpturing just bump at the bottom of the puncture. Minute anterior of the basal bead; this sculpturing is punctures are generally not seen with 12.5 x referred to as the band of punctures, rugose magnification, but are easily seen with 50 x band, etc. Pronotal sculpturing tends to de­ magnification. Small punctures are easily seen crease proportionately as the size and de­ with 12.5 x magnification and can be seen velopment of the horns increases and vice with the naked eye. Large punctures are easily versa. seen without the aid of instruments. A small Horns: Most of the species possessing horns are puncture is termed deep if the bottom of the divided into "majors" and "minors" for de­ puncture cannot be easily seen, and is deemed scriptive purposes. Majors are those individu- shallow if the bottom can be readily observ\ 98 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Large punctures are usually shallow, but may ward projecting horn in males of many species, be termed deep if they are roughly V4 mm or horn variable in length and structure; in addition, more in depth. males of many species possess 2 triangular bos­ R. ses or horns postero-Iateral of anterior horn, Genus STATEGUS Kirby horns variable in length and structure. Meso­ sternum: Completely and setigerously punctate STRATEGUS Kirby in Kirby and Spence 1828: or anterior half setigerously punctate. Elytra: 349 (not Kirby and Spence, loc. cit., as re­ Scutellum large, triangular, basal V~1.3 variably ported by Neave, 1940). [Type species: punctate to rugose. Sutural stria usually a dis­ Strategus aloeus (L.) 1758: 345; subsequent tinctly impressed line, although nearly obsolete designation by Hope (1837: 87), not Casey in one species and punctate in another. Disc 01 (1915: 238) nor Paulian (1947: 78).] elytra variably punctate; punctures minute to subgenus STRATEGODES Casey 1915: 245. large, simple to ocellate to ocellate-umbilicate, [Type species: here designated as Strategus shallow to deep; striae, if present, feebly impres­ antaeus (Drury) 1773: 74. Casey did not desig­ sed, incomplete; sides as disc, humeral umbone nate a type for the subgenus nor have sub­ prominent. Legs: Anterior tibia usually quad­ sequent authors.] ridentate, tarsus almost as long as tibia. Middle . subgenus STRATEGOPSIS Chapin 1932a: 302. and posterior tibiae each with 2 strong, oblique, [Type species: Strategus sarpedon (Burm.) spinose ridges; apices variably toothed or shar­ 1847: 122.] ply angulate. Abdomen: Propygidium with ANASTRATEGUS Casey 1915: 231. [Type numerous fine to moderately coarse, transverse species: Strategus cessus LeConte 1866: 382.] stridulatory ridges. Pygidium in lateral view en­ tirely convex to convex basally and variably con­ Description.-Scarabaeidae, Dynastinae, cave apically, occasionally protuberant at mid­ Oryctini. Form: Oblong-ovate, robust to very dle; basal margin always very coarsely and robust, sides subparallel, dorsum convex, apical setigerously punctate to rugo-punctate in a wide, half of elytra broadly rounded; length from apex transverse band; disc and apical margins vari­ of clypeus to apex of elytra 19.5-80.0 mm. Color: ably sculptured and frequently setigerous; lat­ Castaneous, piceous or black, highly polished to eral emarginations on apical margins either side dull, not shining; vesiture normally ferruginous. of middle present or not in females. Apex of last Head: Shape subtriangular; 2 conical tubercles visible sternite emarginate to subtruncate in delineate front from clypeus, tubercles obsolete males, entire in females. to strong, distinctly separated to connected by a Diagnosis.-Strategus may be distinguished transverse carina. Eyes large; interocular width from other genera in the tribe Oryctini by the 1.66-4.5 transverse eye diameters. Antennae 10 following combination of characters: apex of segmented, club with 3 segments. Clypeus hind tibiae with sharp angulations or teeth, man­ trapezoidal to triangular, sides sinuate, at­ dibles exposed and tridentate, pronotum usually tenuate; apex reflexed, entire to excised; lateral foveate and at least tuberculate subapically. margin of disc usually carinate with sides weakly to sharply declivous, disc variably sculptured. Range.-The combined distributions of the Mentum with disc variably punctate, setigerous, known species extends from the northeastern apex entire to emarginate. Mandibles usually United States, south to Argentina, and includes large, always exposed, tridentate. Pronotum: the Greater and Lesser Antilles. Widest at about middle, apex about 2h as wide as base, margins distinctly beaded, surface variably KEY TO THE SPECIES OF STRATEGUS sculptured, basal margin just inside of bead with or without a transverse, punctate to rugose band 1. Apex of last visible sternite emarginate of sculpturing. Anterior half usually clearly to subtrunctate (Males) ...... 2 foveate, fovea shallow to deep. Subapical margin Apex of last visible sternite entire tuberculate in females, tuberculate or with a for- (Females) ...... 36 A REVISION OF THE GENUS Strategus I 99

MALES Venezuela ...... surinamensis surinamensis Burm. 2(1 ). Mesosternum completely, setigerously 8(4). Apex of clypeus acutely pointed. Middle punctate ...... 3 lobe of mandible very long and slen­ Mesosternum with at least posterior 1/4 der, apex acutely pOinted (Fig. 71). to 1/3 without setigerous punctures 9 Base of pronotum with a very narrow 3(2). Disc of elytra with distinct rows of band of pu nctu res. Southcentral punctures. San Salvador Island, United States ...... mormon Burm. Bahamas ..... : .... .at/anticus n. sp. Apex of clypeus narrowly truncate. Disc of elytra without distinct rows of Middle lobe of mandible short and punctures ...... 4 broad. Base of pronotum without 4(3). 2 to 3 short rows of moderate to large, sculpturing. Grenada (Lesser ocellate and/or umbilicate punctures Antilles) ...... tarquinius n. sp. behind humerus ...... 5 9(2). Mandibles in dorsal view nearly square Rows of ocellate and/or umbilicate (Fig. 44). Pronotum never with horns, punctures behind humerous lack- but with a moderate to strong tuber­ ing ...... 8 cle on midapex. Disc of mentum usu­ 5(4). Apex of anterior horn usually slightly ally coarsely punctate; punctures expanded and weakly to moderately large, dense, often confluent. South­ excised (Fig. 50). Colombia ...... west United States to northwest ...... fascinus Burm. Mexico ...... cessus LeC. Apex of anterior horn entire ...... 6 Mandibles in dorsal view not square. 6(5). Apex of elytra with minute setae (when Pronotum arr(led or not. Disc of men­ not worn off). Posterior horns in tu m not coarsely pu nctate as majors long, slender, forward project­ above ...... 10 ing, and with moderately large 10(9). Disc of elytra, at least on lateral third, punctures apically. Northwest South with definite rows of moderate to America to Mexico. jugurtha Burm. large ocellate and/or umbilicate Apex of elytra without setae. Poste­ punctures ...... 11 rior horns of majors short, stout, usu­ Disc of elytra without definite rows or ally suberect, and with small moderate to large ocellate and/or punctures apically...... 7 umbilicate punctures ...... 21 7(6). Base ot, pronotum with rugose band 11 (1 0). Sutural stria a row of close set, moder­ narrow to moderate. Punctures on ate to large, umbilicate punctures, disc of pronotum small. Anterior horn punctures not forming an impressed of majors with moderate sized line. Guadeloupe .... .syphax (Fabr.) punctures. Punctures on disc of Sutural stria an impressed line, elytra small. Pygidium usually with a punctate or not within; if punctate, small, medio-apical band of setiger­ punctures small, often confluent, not ous punctures. Amazon River south umbilicate ...... 12 to Argentina ...... 12(11). Apex of clypeus moderately to strongly ...... surinamensis hirtus Sternb. excised. Length greater than 50 mm. Base of pronotum with rugose band South America .... .centaurus Kolbe obsolete to narrow. Punctures on Apex of clypeus rounded to truncate or disc of pronotum very small. Anterior weakly emarginate. Length less than horn of majors with small punctures. 50 mm ...... 13 Punctures on disc of elytra minute to 13(12). Elytra shining. Base of pronotum with a very small. Pygidium without a moderate to wide rugose or punctate med io-apical band of setigerous band. Anterior horn, if present, al- punctures. Amazon River north to ways with apex entire ...... 14 100 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Elytra dull, not shining. Base of pro­ bosses. Fovea of pronotum pres- notum with a narrow to obsolete ent ...... 19 rugose band. Anterior horn of all but 19(18). Posterior horns long and slender. Apex the smallest specimens always with of anterior horn expanded and mod­ apex forked, excised, or emar- erately to deeply forked (Fig. 79). ginate ...... 17 Jamaica ...... simson (L.) 14(13). Length less than 30 mm. Pronotum Posterior horns in form of low, pyrami­ never with horns. Cuba ...... dal bosses. Apex of anterior horn not ...... sarpedon (Burm.) expanded or forked, but with a small Length greater than 30 mm. Pronotum triangular notch ...... 20 armed or not ...... 15 20(19). Paramere of genitalia with a spiniform 15(14). Pronotum without posterior horns. process on lateral margin near apex Mandibular lobes confluent, not dis­ (Figs. 110-111). Hispaniola ...... tinct. Disc of pygidium setigerously · ...... aenobarbus (Fabr.) punctate in a broad, irregular band at Paramere of genitalia lacking a midline. Haiti ...... inermis Arrow spiniform process on lateral margin Pronotum with 3 horns except in near apex (Figs. 112-113). Cuba .... minors, and then posterior horns re­ · ...... ajax (Oliv.) duced to triangular bosses. Mandibu­ 21(10). Sutural stria effaced, although occa­ lar lobes usually distinct. Dise of sionally prES ent near apex. Eastern pygidium not setigerously punctate at United States ...... antaeus (Drury) midline ...... 16 Sutl· ral stria present ...... 22 16(15). Disc of elytra adjacent to sutural stria 22(21). Anterior horn with apex expanded and with moderate to large ocellate­ forked, less so in minors where apex umbilicate punctures (Fig. 93). still clearly notched and not merely Cuba ...... symphenax n. sp. weakly emarginate. Paramere of Disc of elytra adjacent to sutural stria genitalia with a well-developed, angu­ without moderate to large ocellate- lar prominance on lateral margin near umbilicate punctures. Cuba ...... apex. Elytra dull, not shining ..... 23 ...... anachoreta Burm. Anterior horn with apex attenuate and 17(13). Paramere of genitalia weakly flared on entire to weakly emarginate. Para­ lateral margin near apex (Figs. 166- mere of genitalia lacking any angular 167), never with a well-developed an­ prominance on lateral margin near gular prominence. Anterior horn in apex. Elytra shining ...... 24 majors distinctly expanded and 23(22). Interocular width about 1 213 transverse forked apically (Fig. 95). Puerto Rico, eye diameters, never more than 2 Virgin Islands ...... talpa (Fabr.) diameters. Posterior horns com­ Paramere of genitalia with a well­ pletely lacking. Cayman Islands ..... developed angular prominence on · ...... caymani n. sp. lateral margin near apex. Anterior Interocular width 2 or more transverse horn in majors expanded or not .. 18 eye diameters. Posterior horns pres­ 18(17). Interocular width about 1 213 transverse ent in majors, minors with at least low eye diameters, never more than 2 bosses. Jamaica ...... simson (L.) diameters. Posterior horns com­ 24(22). Elytra behind humerus with 2-3 short, pletely lacking. Fovea of pronotum distinct rows of moderate to large nearly obsolete. Cayman Islands .... ocellate and/or umbilicate punct- ...... caymani n. sp. ures ...... 25 Interocular width 2 or more transverse Elytra behind humerus without distinct eye diameters. Posterior horns pres­ rows of moderate to large ocellate ent in majors, minors with at least low and/or umbilicate punctures ..... 26 A REVISION OF THE GENUS Strategus / 101

25(24). Apex of clypeus truncate to subtrun­ erately dense punctures. Tubercles cate, occasionally weakly emargi­ on head joined by a strong carina. nate. Middle lobe of mandible short Mexico ...... craigi n. sp. to moderate in length. Brazil to 31 (30). Base of pronotum with rugose band Argentina ...... va/idus (Fabr.) narrow. Sutural stria very feebly im­ Apex of clypeus deeply excised. Middle pressed. Punctures on elytra small. lobe of mandible large and long (Figs. Color black. Mexico...... 63, 65). Central America ...... " ...... , .ado/escens Kolbe ...... Iongichomperus n. sp. Base of pronotum with rugose band 26(24). Apex of clypeus acuminate. Posterior moderate to wide. Sutural stria horns absent. Middle and apical strongly impressed. Punctures of lobes of mandible triangular, both elytra small to moderate. Color pice- prominent, middle large, about twice ous. Mexico ...... howdeni n. sp. as large as apical lobe. Mexico ..... 32(29). Paramere of genitalia roughly triangu­ ...... fallaciosus Kolbe lar, usually distinctly wider at base Apex of clypeus acuminate or not. Pos­ and tapering apically (Figs. 114-117). terior horns present or not. Apical Southern United States to lobe of mandible always much smal­ Brazil ...... a/oeus (L.) ler than middle lobe, never prominent Paramere not as above ...... 33 and triangular ...... 27 33(32). Disc of elytra with punctures small and 27(26). Base of pronotum with rugose band moderate mixed and of moderate narrow ...... 28 density. Interocular width 2213 trans­ Base of pronotum with rugose band verse eye diameters or more. Argen­ wide ...... 29 tina, Uruguay .... .argentinus Kolbe. 28(27). Length less than 35 mm. Posterior Disc of elytra with punctures small and horns absent. Elytra with small minute mixed and sparse. Interocular punctures only. Eastern United width 2 1/2 transverse eye diameters or States ...... sp/endens (Beauv.) less ...... 34 Length greater than 35 mm. Posterior 34(33). Apex of clypeus broadly and shallowly horns present in majors. Elytra with emarginate. Posterior horns short small and minute punctures mixed. and triangular in majors. Base of pro­ Hispaniola, Puerto Rico, Grand notum with rugose band wide. Caymal1 Island .. .ob/ongus (Beauv.) Mexico to Colombia ...... 29(27). Clypeus rounded to subtruncate. Pro­ ...... hipposiderus n. sp. notum never with projecting, acumi­ Apex of clypeus broadly truncate (oc­ nate posterior horns, with rounded casionally very weakly emarginate). bosses at most ...... 30 Posterior horns long and slender in Clypeus emarginate to broadly trun­ majors. Base of pronotum with cate. Pronotum armed with short to rugose band narrow to moderate .,. long, acuminate posterior horns in ...... 35 majors, rounded bosses in minors .. 35(34). Dorsal surface of anterior horn nearly ...... 32 flat. Posterior horns in majors project 30(29). Pronotum with a short anterior horn at about 45° from plane of disc. Mid­ and rounded posterior bosses. Disc dle lobe of mandible 2-3 times larger of pronotum sparsely punctate. than basal lobe. Hispaniola, Puerto Tubercles on head separate to feebly Rico, Grand Cayman Island ...... joined ...... 31 ...... ob/ongus (Beauv.) Pronotum with an anterior tubercle Dorsal surface of anterior horn cari­ only. Posterior armature completely nate. Posterior horns in majors proj­ lacking. Disc of pronotum with mod- ect at about 70° from plane of disc. 102 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Middle lobe of mandible 4-5 times pronotum with a very narrow band of larger than basal lobe. Argentina, punctures. Southcentral United Brazil, Paraguay ...... States ...... mormon Burm...... mandibu/aris Kolbe Apex of clypeus narrowly truncate. Middle lobe of mandible short and FEMALES broad. Base of pronotum without sculpturing. Grenada (Lesser (Females of S. at/anticus, S. craigi, S. fascinus, S. Antilles) ...... tarquinius n. sp. inermis and S. symphenax are unknown. S. at­ 42(36). Mandibles in dorsal view nearly square. /anticus is keyed because I believe the 2 charac­ Disc of mentum usually coarsely ters involved are sufficiently predictive.) punctate, punctures large, dense, 36(1). Mesosternum completely, setigerously often confluent. Southwest United punctate ...... 37 States to northwest Mexico ...... Mesosternum with at least posterior Y4 ...... ~essus LeC. to 113 without setigerous punctures. Mandibles in dorsal view not square...... 42 Disc of mentum usually not coarsely 37(36). Disc of elytra with rows of punctures. punctate ...... 43 San Salvador Island, Bahamas ...... 43(42). Disc of elytra, at least on lateral third, · ...... at/anticus n. sp. with definite rows of moderate to Disc of elytra without rows of large ocellate and/or umbilicate punctures ...... 38 punctures ...... 44 38(37). Elytra behind humerus with 2-3 short Disc of elytra without rows of moderate rows of moderate to large ocellate to large ocellate and/or umbilicate and/or umbilicate punctures ...... 39 punctures ...... 51 Elytra behind humerus without rows of 44(43). Sutural stria a row of close set, moder­ moderate to large ocellate and/or ate to large, umbilicate punctures, umbilicate punctures ...... 41 punctures not forming an impressed 39(38). Apex of elytra with minute setae (when line. Guadeloupe .... .syphax (Fabr.) not worn off). Pygidium nearly im­ Sutural stria an impressed line, punctate to moderately punctate. punctate or not within; if punctate, Northwest South America to punctures small, often confluent, not Mexico ...... jugurtha Burm. umbilicate ...... 45 Apex of elytra without setae. Pygidium 45(44). Length greater than 50 mm. Base of usually densely punctate to sub- pronotum with rugose band wide. rugose ...... 40 South America .... .centaurus Kolbe 40(39). Pygidium setigerously rugopunctate to Length less than 50 mm. Base of pro- subrugose, setae usually dense (ex­ notum with rugose to rugopunctate cept in worn specimens). Disc of pro­ band narrow to obsolete ...... 46 notum with punctures small. Amazon 46(45). Elytra shining ...... 47 River south to Argentina ...... Elytra dull, not shining ...... 48 · ...... surinamensis hirtus Sternb. 47(46). Length less than 30 mm. Pygidium Pygidium setigerously punctate to punctate. Disc of elytra adjacent to rugopunctate, setae sprase to mod­ sutural stria with at least some mod­ erate. Disc of pronotum with erate to large ocellate and/or umbili- punctures very small. Amazon River cate punctures. Cuba ...... north to Venezuela ...... sarpedon (Burm.) · . .surinamensis surinamensis Burm. Length greater than 30 mm. Pygidium 41 (38). Apex of clypeus acutely pointed. Middle rugose. Disc of elytra adjacent to lobe of mandible very long and slen­ sutural stria without moderate to der, apex acutely pointed. Base of large ocellate and/or umbilicate A REVISION OF THE GENUS Strategus / 103

punctures. Cuba ...... punctures ...... 55 ...... anachoreta Burm. 55(54). Elytra behind humerus with 1-4 short to 48(46). Interocular width about 1 213 transverse moderate rows of moderate to large eye diameters, never more than 2 ocellate and/or umbilicate punct- diameters. Cayman Islands ...... ures ...... 56 · ...... caymani n. sp. Elytra behind humerus without distinct Interocular width 2 or more transverse rows of moderate to large ocellate eye diameters ...... 49 and/or umbilicate punctures ..... 58 49(48). Lateral half of disc of elytra with 2 to 56(55). Pronotum with basal band of rugosity several distinct rows of moderate to narrow. Middle and apical lobes of large ocellate and/or umbilicate mandibles subequal. Amazon River punctures ...... 50 south to Argentina .. .validus (Fabr.) Lateral half of disc of elytra usually Pronotum with basal band of rugosity without distinct rows of moderate to wide. Middle lobe of mandible at least large ocellate and/or umbilicate twice as large as apical lobe ..... 57 punctures; if present, feeble and con- 57(56). Pygidium completely rugose and den­ fused. Jamaica ...... simson (L.) sely, setigerously punctate. Interocu­ 50(49). Pygidium not strongly protuberant at lar width greater than 21/2 transverse middle, not overhanging apical mar­ eye diameters. Argentina, Brazil, gin in lateral view. Puerto Rico, Virgin Paraguay ...... Islands ...... talpa (Fabr.) ...... mandibularis Kolbe Pygidium strongly protuberant at mid­ Pygidium moderately rugopunctate to dle, overhanging apical margin in lat- rugose and sparsely setigerous. In­ eral view. Cuba, Hispaniola ...... : .. terocular width less than 2 1f4 trans­ ..aneobarbus (Fabr.) and ajax (Oliv.) verse eye diameters. Mexico to Col- 51 (43). Sutural stria effaced, although occa­ ombia ...... hipposiderus n. sp. sionally present near apex. Eastern 58(55). Interocular width greater than 31/2 United States ...... antaeus (Drury) transverse eye diamters ...... 59 Sutural stria present ...... 52 Interocular width less than 3 transverse 52(51). Elytra dull, not shining. Pronotum with eye diameters ...... 60 basal band of rugosity narrow to ob- 59(58). Color black. Sutural stria very weakly solete ...... 53 impressed. Interocular width greater Elytra sAining. Pronotum with basal than 41f4 transverse eye diameters. band of rugosity variable ...... 54 Mexico ...... adolescens Kolbe 53(52). Interocular width about 1 213 transverse Color castaneous. Sutural stria strongly eye diameters, never more than 2 impressed. Interocular width less diameters. Cayman Islands ...... than 4 transverse eye diameters. · ...... caymani n. sp. Eastern United States ...... Interocular width 2 or more transverse ...... splendens (Beauv.) eye diameters. Jamaica ...... 60(58). Base of pronotum with rugose band · ...... simson (L.) very narrow to obsolete. Pronotum 54(52). Apex of clypeus distinctly (though oc­ usually without a distinct, depressed, casionally weakly) excised. Elytra be­ rugopunctate to rugose patch just hind humerus usually with 2-3 short postero-Iateral of fovea. Hispaniola, rows of moderate to large ocellate Puerto Rico, Grand Cayman Island. and/or umbilicate punctures. Central ...... oblongus (Beauv.) America ...... Iongichomperus n. sp. Base of pronotum with rugose band Apex of clypeus rounded or truncate. moderate to wide. Pronotum usually Elytra behind humerus with or with­ with a distinct, depressed, rugo­ out rows of ocellate and/or umbilicate punctate to rugose patch postero- 104 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

lateral of fovea ...... 61 third. Anterior third rugose. Horns: Anterior 61 (60). Middle lobe of mandible subequal to or short, conical, stout, apex minutely emarginate, slightly larger than apical lobe. anterior face with a faint apical sulcus. Posterior Argentina, Uruguay ...... horns are low, rounded, pyramidal bosses; in ...... argentinus Kolbe lateral view anterior edge slopes forward and Middle lobe of mandible 2 or more downward and dorsal edge is subhorizontal; in times larger than apical lobe ..... 62 dorsal view horns are subparallel, bases joined 62(61). Middle lobe of mandible large, acute. across disc in an arc. Elytra: Sutural stria very Mexico ...... fallaciosus Kolbe weakly impressed, weakly crenulate. Disc acicu­ Middle lobe of mandible moderate in late, sparsely punctate, punctures small, shal­ size, rounded, never acute ...... 63 low; lateral half of disc with 3 very feebly impres­ 63(62). Tubercles on head usually distinctly sed, incomplete striae, striae minutely rugulose separated. Sides of clypeus in dorsal within. Sides weakly wrinkled behind humerus, . view weakly sinuate. Apex of clypeus otherwise as disc. Apex densely punctate, narrowly subtruncate to truncate. punctures small to moderately large. Pygidium: Southern United States to Bra- Convex in lateral view. Disc aciculate, very finely zil ...... aloeus (L.) subgranulate, sparsely punctate, punctures Tubercles on head joined by a moder­ small to moderately large, shallow. Genitalia: ate to strong carina. Sides of clypeus Figs. 108-109. in dorsal view strongly sinuate. Apex Female.-Length 30.5-32.5 mm; width across of clypeus very narrowly subtruncate. humerus 15.0-15.8 mm. As male except in the Mexico ...... howdeni n. sp. following respects: Pronotum: Sides postero­ lateral of fovea with a slightly depressed, Strategus ado/escens Kolbe rounded patch of moderately dense punctures; (Figs. 10-11, 108-109) punctures moderate to large, shallow, and joined to anterior rugosity of pronotum; margin with a Strategus adolescens Kolbe, 1906: 15. [Lec­ band of sparse, moderate to large, shallow totype male (Figs. 10-11), labeled "Mexico punctures. Fovea shallow, longitudinal, hoppe," "TYPE," "27395" and lectoallotype oblong-ovate. Tubercle very low, rounded, nearly female, labeled "TYPE," "27395," here desig­ effaced. Pygidium: In lateral view basal half nated; at ZMHU with my lectotype labels. Also weakly convex, apical half weakly concave. one female para lectotype at ZMHU. Type local­ Lateral depressions shallow. ity: Mexico.] Biology.-Unknown. Male.-Length 31.5 mm; width across Distribution.-Mexico. Considering the fair humerus 16.5 mm. Color black, shining. Head: amount of collecting conducted in Mexico, it Front strongly rugo-punctate. Clypeus with apex might be reasonable to assume a very restricted narrowly truncate, slightly reflexed; surface range (or extinction?) for S. adolescens; this rugo-punctate. Tubercles strong, transverse, might help to account for the lack of additional weakly connected. Mandibles with basal lobe specimens taken during the past 70 years of col­ small, prominently rounded; middle lobe triangu­ lecting. lar, apex rounded; apical lobe similar to middle lobe but smaller. Interocular width 4.5 transverse Locality Records.-3 specimens examined (1 eye diameters. Mesosternum: Anterior half male, 2 females). Specimens deposited at ZMHU. setigerously punctate. Pronotum: Base with a MEXICO (3).-No other data (types). narrow rugose band, band reduced nearly to basal bead at middle. Disc aciculate, sparsely Remarks.-The key characters and the male punctate, punctures small, deep. Sides in basal genitalia should separate this species from all % as disc except for a few moderate, very shal­ others. Variation in characters undoubtedly oc­ low punctures grading to rugosity at anterior curs as it does in all other species in the genus, A REVISION OF THE GENUS Strategus / 105

but it cannot be described here because of the minors). Anterior third of pronotum as disc in lack of additional specimens. majors varying to increasingly rugo-punctate to feebly rugose in less-developed individuals. Horns: Majors (Figs. 12-13) with anterior long, Strategus aenobarbus (Fabr.) stout, parallel sided; apex triangularly notched; (Figs. 1, 12-14, 110-111) dorsal surface with a weak, longitudinal, median carina, and with a similar carina on each lateral Scarabaeus aenobarbus Fabr., 1775: 10. margin. Posterior horns are strongly developed, [Holotype male at Glasgow University, Glas­ laterally compressed pyramidal bosses; in lateral gow, Scotland. Type locality: America, as orig­ view anterior edge vertical, dorsal edge slopes inally given by Fabricius, but here restricted to forward and slightly downward; in dorsal view Dominican Republic.] horns subparallel, bases joined across disc in an Scarabaeus eurytus Fabr., 1775: 7. [Holotype arc, arc slightly produced at middle. Minors (Fig. :male at Glasgow University, Glasgow, Scot­ 14) with anterior tuberculate; tubercle conical, land. Type locality: America, as originally given moderate to large; apex weakly emarginate to by Fabricius, but here restricted to Dominican truncate. Posterior horns red uced to low, Republic.] rounded bosses; bases joined across disc in an Sc.arabaeus aenoburbns Fabr., 1787: 6. [Mis­ arc. Elytra: Sutural stria impressed, crenulate. print. Redescription of S. aenobarbus.] Disc finely granulate, sparsely punctate; Scarabaeus eurytus Fabr., 1787: 5. [Redescrip­ punctures small and minute mixed, shallow; lat­ tion of S. eurytus.] eral 2h of disc with 5-7 regular to irregular rows Scarabaeus fossula Beauvois, 1819: 210. [Types of ocellate-umbilicate punctures; punctures unknown to me, possibly lost. Type locality: small to moderately large, shallow. Sides similar Santo Domingo, Dominican Republic.] New to disc, rows of punctures usually confused. synonymy. Apex moderately densely punctate; punctures Strategus laterispinus Chapin, 1932b: 454. small, shallow. Pygidium: Convex in lateral view. [Holotype male at AMNH. Also one male and Disc finely granulate, sparsely punctate in four female paratypes at AMNH; two male and majors, moderately densely punctate in minors; five female paratypes at USNM. Type locality: punctures small, shallow. Genitalia: Figs. 110- Manville, HaitL] 111. Female.-Length 32.0-38.0 mm; width across Male.-Length 28.8-36.0 mm; width across humerus 14.5-17.6 mm. As male except in the humerus 13.5-17·.2 mm. Color piceous, dull, not following respects: Head: Mandibles slightly shining. Head: Front coarsely rugo-punctate. smaller, especially middle lobe. Interocular width Clypeus with apex truncate, occasionally very 2.66-4.0 transverse eye diameters. Pronotum: feebly emarginate, slightly reflexed; surface Sides just postero-Iateral of fovea usually with a rugo-punctate. Tubercles conical, transverse, slightly depressed, rounded patch of simple to very small. Mandibles small, mostly hidden be­ cresent-shaped punctures; punctures small to neath clypeal fringe; basal lobe small, rounded; moderately large, shallow. Anterior third rugose. middle lobe triangular, apex rounded; apical Fovea shallow. Tubercle subconical, very low, lobe similar to middle lobe but smaller. Interocu­ transverse. Pygidium: Strongly protuberant at lar width 2.33-2.66 transverse eye diameters. middle, overhanging apical margin in lateral Mesosternum: Anterior half setigerously view. Disc densely rugo-punctate to rugose. Lat­ punctate. Pronotum: Base with a narrow to obso­ eral emargination deep, rugose. lete rugo-punctate band, frequently replaced by sparse punctures laterally. Disc aciculate, very Biology.-U nknown. finely granulate, sparsely punctate; punctures Distribution.-Hispaniola, Navassa Island. small and minute mixed, shallow. Sides similar to disc, lateral margin frequently with a narrow, Locality Records (Fig. 1 ).-47 specimens punctate to rugo-punctate band (particularly in examined (19 males, 28 females). Specimens 106 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM were seen from the following collections: AMNH, simson and is placed in new synonymy under S. BMNH, CASC, TAMU, USNM. aenobarbus because it was described from the Dominican Republic. DOMINICAN REPUBLIC (36).-DISTRITO NA­ The male genitalia will serve to distinguish this ClONAL (6): Santo Domingo; LA ALTAGRACIA (5): species from S. ajax which is otherwise mor­ La Romana; LA VEGA (4): Constanza, Jarabacoa; phologically identical. I have not been able to PERAVIA (1): no data; PUERTO PLATA (1): Puerto separate the females of the two species except Plata; SAMANA (6): Sanchez; SAN CRIST6BAL (8): by association with the males and by place of San Cristobal; SANTIAGO (2): Los Hidalgos, San­ origin. tiago; No data (3). January (2), FebruarY'(1), March (1), May (5), June (6), July (2), August (1), September (2), November (1), December (6). Strategus ajax (Olivier) HAITI (10).-OUEST (7): Kenscoff, Manville, (Figs. 1, 15-17, 112-113) Petionvifie, Port-au-Prince; No data (3). February (2), September (3), November (2). Scarabaeus ajax Olivier, 1789: 27. [Type proba­ bly lost (Besuchet, personal communication). NAVASSA ISLAND (1).-No data (1). Type locality: unknown, but designated by Remarl

totype male, labeled "Tex," "Casey bequest ously punctate. Pronotum: Base with a wide to 1923," "Type USNM 48633," and "roosevelti very wide rugose band, band reduced at middle Csy.," here designated; at USNM (Casey col­ almost to basal bead. Disc aciculate, sparsely lection) with my lectotype label. Lectoallotype punctate; punctures small, deep; occasionally female, labeled as male except for type label with minute, sparse secondary punctures. Sides which reads "roosevelti-2, Paratype USNM aciculate, a little more densely punctate; 48633," here designated; at USNM with my lec­ punctures small to large; lateral margin fre­ toallotype label. Also one female paralectotype quently with a moderately wide rugo-punctate to at USNM. Type locality: Texas, United States.] rugose band. Base of posterior horns usually New synonymy. with a small, slightly depressed, densely Strategus frontalis Casey, 1915: 243. [Holotype punctate to rugose area; occasionally a band of male at USNM. Type locality: San Pedro Sula, punctures or rugosity extends from base of pos­ Hond,uras.] New synonymy. terior horns obliquely to posterior angle. An­ Strategus tarsalis Casey, 1915: 243. [Holotype terior Y3 to Y2 rugose. Fovea deep, divided long­ f~male at USNM. Type locality: Texas, United itudinally down middle by a broad, low, rounded States.] New synonymy. carina extending posteriorly from base of an­ Strategus gaillardi Casey, 1915: 244. [Holotype terior horn; anterior angles and fovea either side fem~le at USNM. Type locality: Culebra, of median ridge feebly to strongly rugo-punctate Panama.] New synonymy. to rugose, rarely without sculpturing. Horns: Majors (Figs. 18-22) with anterior moderate in Male.-Length 31.0-60.9 mm; width across length, very stout, attenuate, curving forward humerus 13.8-30.0 mm. Color piceous to black, and upward; apex subtruncate to emarginate; shining. Head: Front densely punctate to rugose emargination feeble and shallow to distinct and behind tubercles; punctures large, shallow, usu- . V-shaped; dorsal surface nearly flat to rounded, ally confluent; medially front varies from im­ occasionally weakly subcarinate longitudinally punctate to variably punctate (punctures small to down middle. Posterior horns short to moder­ large) to rugose; setigerous above eye in unworn ately long, very strongly laterally compressed, specimens. Clypeus with apex variable; speci­ variably attenuate, apex acutely rounded to mens north of Panama usually with apex sub­ broadly and obliquely truncate with 1-3 weak truncate and very feebly to moderately broadly lobes; in lateral view horns project forward and emarginate, rarely with a small triangular notch upward at about 25-650 from plane of disc; in at base of emargination; specimens south of dorsal view horns subparallel to curving slightly Panama usually with,apex variably emarginate as toward one another; bases joined across disc in northern form and with notch at base of emargi­ an arc; arc slightly produced at middle. Minors nation small, triangular, and shallow (very rarely (Figs. 23-24) with anterior similar to that of absent) to large, deep, and U-shaped; apex majors except short in length, apex truncate, and moderately to strongly reflexed; surface acicu­ dorsal surface rounded, not carinate. Posterior late, densely punctate (punctures moderate in horns reduced to very short, rounded, laterally size) to rugo-punctate to rugose. Tubercles in compressed horns or low, rounded bosses. specimens north of Panama conical, transverse, Elytra: Sutural stria strongly impressed, weakly low to moderate, distinctly separated to dis­ to strongly crenulate. Disc aciculate, sparsely tinctly but weakly joined by a low, transverse punctate; punctures small (rarely moderate in carina; tubercles in specimens south of Panama size), shallow, and weakly ocellate-umbilicate in as above but usually stronger and distinctly specimens from south of Panama; 1-3 feebly im­ separated. Mandibles with basal lobe small, pressed, incomplete stria on lateral half of disc. rounded to prominently rounded; middle lobe Sides aciculate, wrinkled behind humerus, subtriangular, large to very large, apex broadly sparsely punctate, punctures small to moderate, rounded; apical lobe small, rounded to subtrian­ shallow. Apex moderately punctate, punctures gular. Interocular width 2.0-2.66 transverse eye small, shallow. Pygidium: Weakly convex in lat­ diameters. Mesosternum: Anterior half setiger- eral view. Specimens north of Panama with disc A REVISION OF THE GENUS Strategus / 109 sparsely punctate; punctures minute, very shal­ (Cocos nucifera L.) and Erythrina g/auca Willd. in low; some specimens from Yucatan, Mexico, and Surinam (Van Dinther, 1956). Bodkin (1919) re­ many from south of Panama also with a few ported small colonies of larvae in decaying moderate, very shallow, setigerous punctures at stumps in Guyana, and I have found numerous apex. Genitalia: Figs. 114-117. Specimens north larvae in a rotten log while collecting in Peru. of Panama (Figs. 114-115) usually with para­ Clusters of larvae are a result of a fortuitous meres subslender, attenuate apically, widest just oviposition site and do not indicate a sub-social before middle; specimens south of Panama ethology as is found in many passalids. Van (Figs. 116-117) occasionally as above but more Dinther also observed in Surinam that larvae frequently with parameres very robust, only were never detected near the roots or stem of a slightly attenuate apically. living coconut palmw hereas, on the other hand, Costa Lima (1953) noted that in Brazil larvae at­ Female.-Length 31.5-55.8 mm; width across tack the roots and lower part of the stipe of new humerus 16.0-26.0 mm. As male except in the palms. Larvae have also been taken from mango following respects: Head: Front punctate roots (Mangifera indica L.) (Bodkin, 1919). Under (punctures small to large mixed) to rugo­ most circumstances larvae probably feed exclu­ punctate to rugose. Clypeus with apex narrowly sively on decayed wood although they will appar­ to broadly subtruncate or rounded, feebly to ently feed on root material when necessary. Pu­ moderately reflexed; surface rugose. Mand i bles pation occurs in the food substrate where an similar to those of male but smaller, especially oval pupal chamber is formed. middle lobe. Pronotum: Sides rugo-punctate to Hurpin and Mariau (1966) conducted labora­ rugose. Anterior half rugose. Fovea moderately tory rearing experiments using 219 larvae and deep. Tubercle conical, moderate to large, trans­ 281 adults from Colombia. The larvae were fed verse, apex usually very feebly emarginate on a mixture of poplar wood and dried cow ma­ (rounded in worn individuals). E/ytra: Sides oc­ nure while the adults were maintained best on casionally with sparse, feeble, ocellate­ banana and orange. In the laboratory adult males umbilicate punctures. Pygidium: In lateral view lived up to five months and adult females up to basal half convex, apical half concave. In speci­ six months. The maximum number of eggs laid mens north of Panama disc usually sparsely to by a single female was 42; eggs were deposited moderately punctate; punctures small to moder­ over a period of 1-3 months with several eggs ate, shallow, punctuation occasionally very re­ laid weekly. Duration (maximum?) of each stage duced; apical margin with a broad rugo-punctate was as follows: egg, three weeks; first instar, two to rugose band; apex with a few setigerous weeks; second instar, three weeks; third instar, punctures; lateral'emargination very shallow. seven months; pre-pupa, two weeks; pupa, six Specimens south of Panama as above, but discal weeks; adult, six months. This provided a total punctures usually moderately large and con­ life span of 17 months. Other lengths reported fluent; apical V3 to Y2 completely and densely for pupation are 31 days (Van Dinther, 1956) and setigerously punctate. 43 days (Duges, 1886). Adults can usually be Biology.-Duges (1876, 1886) was the first to found year round where the climate permits. describe the larva and pupa of S. a/oeus, and Adult beetles have been found feeding on Ritcher (1944, 1966) has provided the most re­ roots of date palms (Phoenix dactyliferas L.) in cent and accurate description of the larva. Eggs Arizona (Cockerell, 1906); wax palms (prob. are usually deposited in dead or rotten wood Copernicia cerifera Mart.) in Brazil (Goncalves, where the larvae develop. Larvae have been 1946); young oil palms (E/aeis guineensis Jacq.) found under old logs and boards in Louisiana in Colombia (Vayssiere, 1965); Furcraea cabuya (Schufeldt, 1884), in a rotten ash tree (Fraxinus Trel. and the leaves of Agave picta Salm-Dyck in sp.) in Texas (Ritcher, 1966), from the trunk of a Costa Rica (Nevermann, 1933); and coconut live oak tree (Quercus sp.) in Mexico (Duges, palms (Cocos nucifera L.) in Guyana (Bodkin, 1876,1886), and in sawdust piles at sawmills and 1919), Surinam (Van Dinther, 1956), Mexico beneath the trunks of felled coconut palms (Kolbe, 1906), and Venezuela and Brazil (Lever, 110 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

1969). Label data indicated feeding on oil palms mens examined (751 males, 1,452 females). in Colombia and Ecuador, coconut palms in Specimens were seen from the following collec­ Costa Rica and Venezuela, sugar cane in Col­ tions: AHCC, AMNH, BCRC, BMNH, CASC, CNCI, ombia, and in rotten wood in Texas, Mexico, and DEUN, FMNH, HAHC, INHS, JOG, KSUC, LACM, Peru. MCZC, MHNCM, OSUC, OSUO, PMNH, RLWE, In Van Dinther's (1956) study, feeding damage SBMN, SEMC, TAMU, UC, UCMC, UMMZ, USNM, was restricted to young coconut palms up to four USP, UWEM, UZM, ZMHU. years of age. Reasons were not given as to why BELIZE (5).-STANN CREEK (5): Placentia Feb­ older palms were not attacked. Adult beetles ruary (1), March (4). burrowed down as far as 40 cm near the base of the palm to a position under the stem base. Tun­ BOLIVIA (12).-BENI (10): Guayaramerin, Reyes, neling tt"!en proceeded upward into the heart of Rurrenbaque (Rio Beni), San Joaquin; the wood where a 3-5 cm wide shaft was formed. COCHABOMBA (1): Chapare; No data (1). January Trees so infested wilted and eventually died. (2), April (1), May (5), June (2), November (1), De­ Usually one beetle was found per tree, but as cember (1). many as four of both sexes have been found in a single palm. However, Strategus a/oeus is rarely BRAZIL (38).-AMAZONAS (7): Manaus, Porto of economic significance to palm plantations. Santana, Porto Velho de San Antonio, Taracua, These large beetles apparently have few Tefe; BAHIA (2): No data; CEARA (2): Fortaleza, natural enemies. The larvae have occasionally Villa Hazare; GOlAS (1): Magalhaes; GUANABARA been eaten by aboriginal indians in Guyana (1): Serra da Carioca; MATO GROSSO (1): Amazon (Bodkin, 1919) and undoubtedly in other tropical region; PARA (16): Belem, Maves, Obidos, regions as well. The larvae are also a common Oriximina, Santarem; PERNAMBUCO (1); Recife; intermediate host for Macracanthorhynchus . No data (6). January (1), February (2), March (1), hirudinaceus (Pallas) (Archiacanthocephala: April (1), May (1), July (2), August (9), September Oligacanthorhynchidae) in Brazil (Costa Lima, (1), November (2), December (1). 1953) and possibly in Louisiana (Manter, 1928). COLOMBIA (143).-AMAZONAS (11): Leticia; AN­ The larval host dies after several weeks of infec­ TIOQUIA (1): Antoquia; ATLANTICO (2): Barran­ tion by these parasites. Adults have been ob­ quilla; CHoc6 (2): Frigana; CUNDINAMARCA (3): served trapped in large spider webs in Venezuela Bogota, Fusa(gasuga?); MAGDALENA (28): on two occasions (Beebe, 1944), and all stages in Aracataca, Medialuna, Parque Tayrona (21 mi. E. the life cycle are probably susceptible to various Santa Marta), Rio Frio, Sevilla; META (1): Cano fungi and bacteria. Grande near Villavicencio; NARINO (1): Tumaco; Beebe (1944) has demonstrated in Venezuela SANTANDER (3): Rio Opon, Velez; VALLE DEL that the males will fight with one another, pre­ CAUCA (11): Anchicaya Dam (70 mi. E. Buenaven­ sumably for females, with the advent of the rainy tura), Cali, Palmira, Rio Anchicaya between season or by artifically induced "rain." He stated Buenaventura and Cali; No data (80). February that the rains "unlock the reproductive reac­ (12), March (3), April (7), May (16), June (4), July tions" of these insects. While reproductive activ­ (3), August (2), October (1), November (1), De­ ity is probably initiated by the rains, Beebe's data cember (2). are insufficient to warrant describing the males as fighting "for" the female, especially when COSTA RICA (93).-ALAJUELA (1): Alajuela; CAR­ other studies have shown this not to be true (see TAGO (4): Irazu, Tres Rios Turrialba; GUANACASTE Arrow, 1951). (3): Finca Jimenez (near Taboga), Liberia, Tenorio; HEREDIA (1): no data; LIMON (15): Distribution.-Strategus a/oeus is the most Guapiles, Hamburg Farm (Reventazon), Las widespread species in the genus, occurring from Mercedes, Port Limon, Reventazon; PUNTARENAS the southern United States through Central (23): Bocca de Barranca, Esquinas near Golfito, America to central Brazil and Bolivia. Monteverde de Puntarenas, Osa Productos Locality Records (Figs. 5, 8).-2,203 speci- Forest, 1.8 mi. W. Rincon; SAN JOSE (29): San A REVISION OF THE GENUS Strategus / 111

Jose; No data (17). February (9), March (5), April Yerba Buena (20 mi. N. Bochil), 25 mi. E. (7), May (18), June (13), July (1), August (11), Sep­ Zanatepec, Jct. Rts. 190 & 195; CHIHUAHUA (16): tember (22), October (2), December (1). Buena Vista, Camargo, 20 mi. SW. Camargo, Catarinas, 10 mi. S. Las Delicas, Primavera, San ECUADOR (6).-GUAYAS (2): Balzar, Naranjal; Francisco del Oro, Santa Barbara; COAHUILA PACHINCHA (2): Santo Domingo de 105 Col­ (10): Muralla, Parras de la Fuente, Ramos Arizpe; orados; No data (2). March (1), August (1), De­ COLIMA (1): Manzanillo; DISTRITO FEDERAL (8): cember(2). Escandon, Jacubaua, Mexico City, Nuevo Bos­ EL SALVADOR (21).-LlBERTAD (1): 20 mi. E. La que Chapultepec, Tocubaya; DURANGO (19): 7 Libertad; SAN SALVADOR (16): Lake IIl0pango, mi. SW. Cuencame, Durango, San Juan del Rio, San Salvador, Santo Tomas; SONSONATE (1): San Pedro de Guanace, Tlahualilo; GUANAJUATO Sonsonate; No data (3). March (1), May (14), June (2): 22 mi. E. Penjamo, San Miguel de Allende; (3), ~ugst (1). GUERRERO (3): Iguala, Irapuato, Rio Balsas; FRENCH GUIANA (11).-ININI (7): Maroni River, HIDALGO (5): Guerrero Mill., 22 mi. NE. Jacala, Saul (Gruner, 1971); GUYAN (4): Cayenne, Crique Pachuca; JALISCO (63): Ajijic, Guadalajara; Gregoire (FI. Sinnamary) (Gruner, 1971). October MEXICO (1): Chapingo; MICHOACAN (6): Morelia, and November (9). Patzcuaro, Playa Azul, Tancitaro, Tuxpan; MORELOS (10): Antiguo, Cuernavaca, Progresso; GUATAMALA (36).-ALTA VERAPAZ (5): Baleu, NAYARIT (14): 20 MI. SE. Ixtlan del Rio, Tepic, 24 Coban, Finca San Juan, Senahu; CHIMAL­ mi. SE. Tepic; NUEVO LEON (99): Chipenque TENANGO (2): S. P. Yepocapa; CHIQUIMULA (1): Mesa (Monterrey), Hacienda Vista Hermosa (Villa Chiquimula; GUATAMALA (1): Guatamala City; Santiago), Linares, 28 km. W. Linares, 28 km. IZABAL (2): Puerto Barrios; PETEN (8): Sabaneta, NW. Linares, Monterrey, 5 mi. S. Monterrey, Tikal; SASATEPEQUEZ (3): Acatenango, Santa Rancho Presa, 17 km. N. Sabinas Hidalgo; OAX­ Maria de Jesus; SAN MARCO (6): Ayutla, Puente ACA (50): 2.7 mi. NW. EI Camaron, 11.6 mi. W. Talisman; SOLOLA (1): Panajachel; SUCHI­ Jalapa del Marques, 22 mi. S. Jesus Carranza, 20 TEPEQUEZ (3): Finca Moca, Cuyotenango; ZACAPA mi. S. Juchatenango, Juquila Mixes, 3 mi. E. La (1): Zacapa; No data (3). February (1), March (4), Ventosa, 8 mi. N. La Ventosa, Oaxaca, Ocotlan, April (3), May (5), June (9), July (4), August (2), Salina Cruz, Tehuantepec, 6 mi. W. Tehuan­ November (1). tepec, 12 mi. W. Tehuantepec, Temascal, Tux­ GUYANA (13).-DEMERARA (3): Georgetown; Es­ tepec; PUEBLA (15): 3 mi. W. Acatepec, Acatlan, SEQUIBO (4): Essequibo, Katabo (Bartica), Penal Finca San Juan Apulco (near Zacapoa), 13 mi. N. Settlement (Bartica District); No data (6). De­ Huauchinongo, Tezuitlan, 3 mi. E. Tezuitlan, 6 cember (1). mi. W. Tezuitlan, Xicotepec de Juarez, Zacapoax­ tla; QUINTANA Roo (2): Felipe Carrillo Puerto; HONDURAS (10).-ATLANTIDA (1): Tela; CORTES SAN LUIS POTOSI (106): 18 mi. E. Ciudad Maiz, EI (1): Puerto Cortes; FRANCISCO MORAZAN (5): Esc. Naranjo, EI Saito, EI Saito Falls, Palitla, 2 mi. SE. Ag. Pan. (Zamorana), Teguicigalpa; YORO (3): EI Pedro Montoya, Saltillo, 30 mi. SW. San Luis Progreso, 12 km. W. Olanchito. Potosi, Tamazunchale, 25 mi. N. Tamanzun­ MEXICO (842).-AGUASCALIENTES (6): Aguas­ chale, Valles, 36 km. W. Valles, 3 mi. W. Xilitla; calientes; CAMPECHE (3): Camp~che, SINALOA (30): Excuinapa, Los Mochis, Mazatlan, Matamoros; CHIAPAS (75): Arriga, 7.2 mi. SE. 5 mi. N. Mazatlan, 9 mi. N. Mazatlan, 4 mi. N. San Chiapa de Corzo, 3 mi. SE. Comitan, 5 mi. SW. EI Bias, 27 mi. E. Villa Union.; SONORA (10): Alamos, Bosque, EI Rincon, 25 mi. N. Huixtla, Lagos des 10 mi. NE Ciudad Obregon, Hermosillo, Navajoa, Colores (Rt. 17), 10 mi. S. Malpaso, Ocosingo, 30 mi. E. Ures on road to Moctezuma; TABASCO Ocozocoautla, 12 mi. N. Ocozocoautla, 9 mi. N. (6): Frontera, Ocotlan; TAMAULIPAS (48): Ciudad Ocozocoautla, Palenque, 5 mi. NW. Pijijiapan, Mante, Ciudad Victoria, Gomez Farias (Rio Frio), Puerto Arista, San Cristobal, 6 mi. E. San Cris­ Guemes, Hidalgo," Llera, Tampico; VERACRUZ tobal, 8 mi. NE. San Cristobal, 2 mi. S. Simojovel, (103): 1 mi. N. Anton Lizardo, Banderilla, Tonola, Tres Picos (Hwy. 200), Tuxtla Gutierrez, Catemaco, 5 mi. E. Catemaco, Chicontepec, 112 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Cordoba, Coyame (Lake Catemaco), Dos (31): Mesa City, Phoenix, Tempe; Pima (27): Amates, Jalapa, 5 mi. N. Huatusco, Montepio (8 Baboquivari Mts., Continental, Madera Canyon, mi. N. Sontecomapan), Orizaba, Palmo Sola, 4 Santa Rita Mts., Tucson; Pinal (1): Superior; mi. SW. Panuco, Puente Nacional (34 mi. E. Santa Cruz (55): Douglas, Madera Canyon, Jalapa), San Andres Tuxtla, Tierra Blanca; Nogales, Patagonia, Tubac; Yavapai (6): Pre­ YUCATAN (61): Chichen Itza, Piste, Valladolid, scott, Red Rock Camp, Verde Hot Springs; Insuf­ Xcan; No data (50). March (2), April (8), May ficient or no data (11). ARKANSAS (8): Hempstead (123), June (253), July (185), August (16), Sep­ (6): Hope; Miller (2): Texarkana. FLORIDA (1): Es­ tember (34), October (9), December (1). cambia (1): Pensacola. GEORGIA (3): Chatham (1): Savannah; Jeff Davis (1): Hazelhorst; No data NICARAGUA (23).-GREAT CORN ISLAND (2); LE6N (1): LOUISIANA (144): Acadia (1): Crowley; Caddo (3): Momotombo; MANAGUA (12): Managua City; (5): Shreveport; Jefferson (7): no data; Lincoln ZELA~A (1): Blue Fields; No data (5). January (3), (6): Ruston; Madison (2): Mound, Tallulah; June(11 ). Natchitoches (1): Natchitoches; Orleans (11): PANAMA (93).-CANAL ZONE (63): Ancon, Bal­ New Orleans; Ouachita (1): Monroe; Rapides boa,Barro Colorado Island, Fort Clayton, Gatun, (88): Alexandria; St. Landry (1): Opelousas; St. Madden Dam, Margarita; CHIRIQuf (15): Boquete, Martin (1): Belle River; St. Mary (1): Morgan City; David;'EI Valle de Nubes, EI Volcan Chiriqui, Rov­ St. Tammany (8): Covington; Vermillion (11): ira; COCLIS (2): Penonome; COLON (6): Coconut Gueydan. MISSISSIPPI (29): Adams (3): Natchez; Point; DARIEN (1): Darien; HERRERA (1): Santa Forest (1): Hattiesburg; Grenada (4): Grenada; Maria (EI Real); Los SANTOS (2): Sabana; PANAMA Harrison (17): Biloxi, Gulfport, Long Beach; (2): Panama City; No data (1). January (15); Feb­ Yazoo (3): no data; No data (1). NEW MEXICO (1): ruary (9), March (14), April (6), May (14), June (5), ' Dona Ana (1): Mesilla Pk.; Grant: no data (Fall July (2), September (2), November (2), December and Cockerell, 1907). TEXAS (249): Bastrop (1): Bastrop; Bexar (6): San Antonio; Brazoria (2): (7). Freeport; Brazos (9): Bryan, College Station; PERU (37).-HuANUCO (5): Tingo Maria; LORETO Brewster (16): Alpine, Boquillas Basin, Chisos (26): Iquitos, Pucallpa, Yarina Cocha (Rio Ucay­ Mts. Basin (Big Bend Nat'l. Park), Glenn Spring, ali); PIURA (2): Quiroz (Rio Paucartambo); No 5 mi. N. Glenn Spring, Juniper Canyon (Chisos data (5). January (2), February (12), April (3), May Mts.), Panther Junction (Big Bend Nat'l. Park), (3), June (4), September (4), November (6). Rio Grand Camp (Big Bend Nat'l. Park); Brown (1): Brownwood; Burleson (4): Clay, Somerville; SURINAM (7).-MAROWIJNE (1): Anapaike (Rio Calhoun (2): Ft. Lavaca; Cameron (33): Lawa); SURINAME (6): Lelydorp Bauxite Mine, Brownsville, Harlingen; Colorado (1): Columbus; Paramaribo. January (1), November (2), De­ Culberson (1): Van Horn; Dallas (12): Dallas; EI cember (1). Paso (2): EI Paso; Gillespie (1): no data; Hardin TRINIDAD (60).-MAYARO (2): Mayaro Beach; ST. (1): Beaumont; Harris (4): Houston, Seabrook; GEORGE (15): Arima Valley, Las Cuevas Bay, Hidalgo (28): Donna, Edinburgh, Mission, San Maraval, Morne Bleu, Port-of-Spain, St. Augus­ Juan; Jeff Davis (13): Davis Mts., Ft. Davis, Valen­ tine; ST. PATRICK (34): Icacos; No data (9). tine; Jefferson (1): Port Arthur; Kerr (4): Kerrville; January (34), February (3), March (3), April' (4), Kleburg (2): Kingsville; La Salle (3): Fowlerton; May (4), June (4), November (1). Mata Gorda (1): Palacios; Maverick (3): Eagle Pass; McLennon (1): Waco; Menard (1): Menard; UNITED STATES (617).-ALABAMA (12): Conecuh Nacodoches (1): Nacodoches; Polk (5): (2): no data; Macon (5): Tuskegee; Mobile (4): Livingston; San Jacinto (3): Cold Spring; Travis Mobile; No data (1). ARIZONA (167): Cochise (15): (10): Austin; Uvalde (7): Sabinal, Uvalde; Val Ash Canyon, Carr Canyon, Portal, Ramsey Can­ Verde (50): Del Rio, Lake Walker, Victoria (9): yon, San Bernadino Ranch, Tombstone; Victoria; Wharton (3): EI Campo; Webb (2): Coconino (2): Sedona; Gila (14): Globe, Winkel­ Laredo; Insufficient or no data (39). January (2), man; Graham (5): Gila Valley, Safford; Maricopa March (4), April (6), May (164), June (129), July A REVISION OF THE GENUS Strategus / 113

(137), August (96), September (23), October (4), given by Laporte is apparently confused and is November (4). not traceable. Arrow (1937a) was also perplexed by Laporte's S. aesalus for the same reasons. VENEZUELA (103).-ARAGUAY (6): Maracay, The description of S. aesalus is only very gen­ Rancho Grande; BOLIVAR (28): Moitaco, Suapure eral, but the sizes indicated by Laporte suggest (Caura River); CARABOBO (7): San Esteban (near that this species should fall into synonymy with Puerto Cabello), Zapateral; DISTRITO FEDERAL S. aloeus and not with S. surinamensis (11): Caracas, EI Limon, La Guaira; FALCON (6): surinamensis which occurs sympatrically at the Boca de Orda, 60 mi. SE. Maracaibo; MIRANDA type locality. (1): Rio Chico; MONAGAS (22): Caripito; SUCRE S. semiramis (Fabr.) and S. piosomus Kolbe (2): Puerto Hierro; TRUJILLO (1): Valera; YARACUY are entered in new synonomy because the types (1): Urama; ZULIA (4): Kasmera (12.5 mi. SW. are conspecific with S. aloeus. Machique), Los Encentados (Rio Onia E. of A clinal mode in character states is to be seen Zulia); No data (14). February (3), March (7), April in a north-south direction. Southern specimens (21), May (12), June (5), July (3), August (1), Oc­ are usually progressively larger, darker, with a tober (15), November (8), December (2). deeper clypeal excision, stouter genitalia in the males (Figs. 114-117), and with increased pygi­ Remarks.-The males of S. aloeus can be dial hairiness in the females. The clinal variation separated from all other species by the charac­ is gradual and not stepped, thus eliminating for teristic shape of the genitalia; they may be easily the present the possibility of subspecies desig­ confused with other species if external features nations. It should be noted that "northern-Ii ke" only are used in identification. The key charac­ individuals (smaller in size, lighter in color, etc.) ters will serve adequately to separate the females occur occasionally in South America, and from all other species. "southern-like" specimens (larger in size, darker Strategus aloeus is the most widespread, in color, etc.) occur occasionally in Central abundant, and morphologically variable species America and Mexico. The Isthmus of Panama in the genus. Until this revision, S. aloeus and S. appears to be a convenient and fairly accurate julian us were considered distinct species, S. locale by which to separate the northern and aloeus occurring primarily in northern South southern populations, but it is by no means a America and S. julianus occurring from the barrier, and mixing between the two populations southern United States to Panama. All previous probably occurs freely. authors have overlooked the use of the male Schaeffer (1915) described a new variety of S. genitalia as a taxonomic character (except ju/ianus from Arizona (var. arizonicus) in which Saylor, 1946) and have not examined as many the males have the posterior pronotal horns specimens as was done here (2,203 examples). pointed (Fig. 20) instead of broadly truncate; this Consequently, there was understandable confu­ character state is part of the normal variation sion as to the separation of these two supposed within the species and is to be seen in other species (see Bates, 1886-1890; Burmeister, localities as well, i. e., Mexico, Central and South 1847; Casey, 1915; Endrodi, 1959; Kolbe, 1906). America. I have seen a number of specimens An analysis of all the taxonomic characters used from NW. Peru which are proportionately some­ in this study revealed that S. julianus was what longer in body length than is considered synonymous with S. aloeus, and so S. ju/ianus normal and also a series from eastern Yucatan in and all its junior synonyms fall into new Mexico which are consistently darker (black) and synonymy under S. aloeus. more highly polished. Individuals from Ven­ Laporte (1840) described S. aesalus as new al­ ezuela and northern Colombia are usually con­ though he may not have meant to do so as he sistently larger than in any other area. It is not cites Fabricius (1801) as a previous (name?) au­ known whether these local variants, or perhaps thor. Fabricius described Aesalus as a new even the entire cline, are a result of genus in the Lucanidae but not as a species of ecophenotypy or whether there are some actual Scarabaeus. In addition, the Fabrician reference differences in gene pools. 114 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Strategus anachoreta Burmeister and slightly forward; in dorsal view horns sub­ (Figs. 2, 25-28, 118-119) parallel, bases joined across disc in a shallow arc. Elytra: Sutural stria impressed, weakly to Strategus anachoreta Burmeister, 1847: 134. strongly crenulate. Disc aciculate, sparsely [Lectotype male and lectoallotype female de­ punctate, punctures small and minute mixed, signated by Endrodi (1973a); these and one shallow; 5-7 rows of ocellate-umbilicate male para lectotype at Martin Luther Univer­ punctures on lateral half, punctures large, shal­ sitat, Halle, German Democratic Republic. low, rows 3 and 6-7 confused. Sides as disc ex­ Type locality: Cuba.] cept with 5 similar rows of ocellate-umbilicate punctures, row 3 confused. Apex densely Male.-Length 35.0--50.0 mm; width across punctate; punctures small to moderately large, humerus 18.0--25.0 mm. Color castaneous to shallow. Pygidium: Convex in lateral view, apical piceoys, shining. Head: Front finely rugose; Y4 frequently concave. Disc very finely granulate, setigerous above eye in unworn specimens. moderately densely punctate, punctures small Clypeus with apex broadly truncate, Slightly re­ and minute mixed, occasionally moderate in flexed; surface finely rugo-punctate. Tubercles size, shallow. Apical margin with a wide, rugose conical, low, occasionally transverse and weakly band; rugosity usually spreading to disc in conne9ted. Mandibles with basal lobe small, minors; occasionally a short, medio-apical row . prominently rounded; middle lobe subtriangular, of setigerous punctures in unworn specimens . . large, apex rounded; apical lobe similar to mid­ Genitalia: Figs. 118-119. dle lobe but smaller. Interocular width 2.5-3.0 transverse eye diameters. Mesosternum: An­ Female.-Length 32.7-42.8 mm; width across terior half setigerously punctate. Pronotum: humerus 14.9-24.0 mm. As male except in the Base with a wide, rugose band; band reduced following respects: Head: Front rugose to rugo­ medially almost to basal bead. Disc aciculate, punctate. Clypeus with apex usually not so punctate; punctures sparse, small, shallow. broadly truncate as male; surface rugose to Sides similar to disc plus the following: lateral rugo-punctate. Mandibles similar but smaller, margin with or without a narrow rugo-punctate especially middle lobe which is nearly subequal band, majors rugose to rugo-punctate from base with basal lobe. Pronotum: Sides just postero­ of posterior horns to anterior angle and into lateral of fovea usually with a slightly depressed, fovea; minors completely rugose in anterior half rounded patch of simple to crescent-shaped of pronotum. Fovea deep, divided in half by a punctures, punctures small to moderately large, low, broadly rounded, longitudinal carina ex­ shallow. Anterior half rugose. Fovea shallow. tending posteriorly from base of anterior horn. Tubercle conical, transverse, very low. Pygidium: Horns: Majors (Figs. 25-26) with anterior moder­ In lateral view basal half convex, apical half con­ ate to long, stout, attenuate, curving forward and cave. Disc entirely rugose to rugo-punctate with upward; apex narrowly rounded, dorsal surface very short, sparse setae in unworn specimens. rounded to nearly flat, occaSionally with a faint, Lateral emargination shallow. longitudinal carina down middle. Posterior horns moderate to long, stout, laterally com­ Biology.-Little is known about the life history pressed, attenuate, apex rounded; in lateral v!ew of S. anachoreta. Label data indicate that it has horns project forward and upward at 45-650 from been taken at lights and that it injures coconut plane of disc; in dorsal view horns parallel, occa­ palms (Cocos nucifera L.). Stahl and sionally with apices turned slightly inwards, Scaramuzza (1929) and Valdes (1951) observed bases joined across disc in an arc. Minors (Figs. that they attack roots and occasionally the seed 27-28) with anterior short, stout, erect, at­ pieces of sugar cane (Saccharum officinarum tenuate, apex narrowly rounded. Posterior horns (L.)) but are not considered major pests. Chapin reduced to low, rounded bosses; in lateral view (1932a) indicated they are an important enemy of dorsal surface subhorizontal to higher at apex, coconut palms. Tiphia argentipes Cresson anterior surface vertical to sloping downward (Hymenoptera: Tiphiidae) and Campsomeris A REVISION OF THE GENUS Strategus / 115 trifasciata (Fabr.) (Hymenoptera: Scoliidae) have paralectotypes also at USNM. Type locality: been listed as natural enemies (Valdes, 1951). Florida, United States.] Strategus pinorum jcuasey, 1915: 248. [Lec­ Distribution.-Cu ba. totype male, labeled "Southern Pines, VII-29, Locality Records (Fig. 2).-147 specimens N.C., A. H. Manee, 07," "Casey bequest 1923," examined (54 males, 93 females). Specimens "Type USNM 48637," and "pinorum CSy.," were seen from the following collections: AMNH, here designated; at USNM (Casey collection) BMNH, CASC, CNCI, MCZC, USNM. with my lectotype label. Lectoallotype female, labeled "Southern Pines, VII-20, N. C., A. H. CUBA (146).-CAMAGUEY (98): Baragua, Jaronu; Manee, 07," "Casey beq uest 1923," HABANA (8): Habana City, La Alianza, Vibora; ISLA "pinorum - 3, Paratype USNM 48637," and DE PINOS (3): Los Indios, Nueva Gerona; LAS VIL­ "pinorum CSy.," here designated; at USNM LAS (16): Bangos de Ciega Montera, Cayamas, with my lectoallotype label. Also one male and Santa Clara, Soledad; ORIENTE (5): Baracoa, two female paralectotypes at USNM. Type loc­ Santiago de Cuba, Sierra de Cristal; PINAR DEL ality: Southern Pines, North Carolina, United RIO (4): Pjnar del Rio; No data (12). January (2), States.] May (43), June (36), July (7), August, (4), Sep­ Strategus septentrionis Casey, 1915: 249. [Lec­ tember (1), October (4). totype male and lectoallotype female here de­ Remarks.--S. anachoreta is easily separated signated: at USNM with my lectotype labels. from all other species by the partially punctate Also three male paralectotypes at USNM. Type mesosternum; the presence of definite rows of locality: New Jersey, United States.] moderate to large, ocellate-umbilicate punctures Strategus sinuatus Casey, 1915: 250. [Holotype on the lateral half of the elytral disc; the broadly male at USNM. Type locality: Alabama, United truncate clypeal apex; the shining elytra; and the States.] presence of three simple horns on the pronotum. Strategus semistriatus Casey, 1915: 250. This species is very closely related to S. sym­ [Holotype male at USNM. Type locality: Texas, phenax which also occurs in Cuba. United States.] Strategus antaeus houstonensis Knaus, 1925: 182. [Holotype male and allotype female pre­ Strategus antaeus (Drury) sumably at KSUC (Knaus Collection). Type (Figs. 4, 29-31, 120-121) locality: Houston, Texas, United States.]

Scarabaeus antaeus Drury, 1773: 74. [Types un­ Male.-Length 18.2-40.7 mm; width across known to me. Type locality: originally pub­ humerus 11.0-19.5 mm. Color castaneous to lished as Jamaica but probably Jamaica, New piceous, shining. Head: Front coarsely rugo­ York, United States.] punctate, setigerous above eye in unworn spec­ Scarabaeus maimon Fabr., 1775: 10. [Holotype at imens. Clypeus with apex rounded, usually with BMNH (Banks Collection) (Zimsen, 1964). Type an acute, subapical tooth in unworn specimens; locality: originally given as America but here surface coarsely rugo-punctate. Tubercles coni­ restricted to eastern United States.] cal, small, connected by a low, transverse carina. Scarabaeus maimon Fabr. 1787: 6. [Redescrip­ Mandibles with basal lobe small, prominently tion.] rounded; middle lobe long, slender, apex very Strategus divergens Casey, 1915: 246. [Holotype narrowly rounded; apical lobe small, rounded. male at USNM. Type locality: Gulf States, Interocular width 2.75-3.0 transverse eye United States.] diameters. Mesosternum: Anterior half setiger­ Strategus atrolucens Casey, 1915: 247. [Lec­ ously punctate. Pronotum: Base with a very nar­ totype male, labeled "Fla.," "Casey bequest row to obsolete feebly punctate to rugo-punctate 1923," "Type USNM 48639," and "atrolucens band. Disc very finely granulate, faintly impres­ CSY.," here designated; at USNM (Casey col­ sed medially, sparsely punctate; punctures lection) with my lectotype label. Two male small, moderately deep. Sides as disc except 116 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM base of posterior horns with an area of simple to coarsely rugose. Mandibles similar to those of crescent-shaped punctures; punctures moder­ male but smaller, especially middle lobe; basal ate to large, dense, shallow. Anterior third as lobe more prominently rounded. Pronotum: disc in majors; minors with anterior angles, Sides with punctures becoming moderate to fovea, base of posterior horns, and sides of an­ large, often crescent-shaped. Anterior Y3-Y2 terior horn rugose; very small individuals with rugose to rugo-punctate, occasionally reduced entire anterior third rugose. Fovea deep, divided (especially in Massachusetts and New Jersey longitudinally by a low, rounded carina extend­ populations). Fovea moderately deep. Tubercle ing posteriorly from base of anterior horn. conical, transverse, low, weakly emarginate. Horns: Majors (Figs 29-30) with anterior very Pygidium: In lateral view basal half convex, api­ long, slender, attenuate, curving forward and cal half feebly concave. Disc moderately to den­ upward, (recurving posteriorly at apex in largest sely setigerously punctate (usually sparsely specimens); apex acutely rounded, dorsal sur­ punctate in Massachusetts and New Jersey face weakly carinate to rounded, usually very populations); punctures large, shallow. Lateral feebly and longitudinally sulcate medially. Poste­ emargination shallow, more densely punctate. rior horns long, slender (but more robust than anterior), laterally compressed, attenuate, apex Biology.-Ritcher (1966) provided a detailed narrowly rounded; in lateral view horns curve description of the third instar larva. Cockerell forward and upward at approximately 45-85° (1906) mentioned that the larvae may injure from plane of disc; in dorsal view horns sub­ peach (Prunus) roots in Alabama, and label data parallel to slightly diverging, apices curving to­ indicate specimens have been taken on peach ward one another; lateral surface convex, me­ roots and trunk, in piles of chicken feathers, and dian surface nearly flat. Minors (Fig. 31) with an­ ,at lights. Manee (1908) observed in the sandy terior short, robust, weakly attenuate; apex usu­ barrens near Southern Pines, North Carolina, ally feebly excised, occasionally blunt. Posterior that S. antaeus dig burrows with a 6-8 inch verti­ horns essentially as in majors except very short cal shaft that end with a 1-5 inch horizontal or reduced to low, rounded to pyramidal bosses; chamber; 2-3 diverging chambers are occasion­ in lateral view dorsal surface subhorizontal, an­ ally present. Each chamber is provided with de­ terior surface subvertical to sloping forward and cayed oak (Quercus) leaves which serve as larval downward; in dorsal view horns subparallel, food. A single egg is placed in each chamber. bases joined across disc in a weak, medially pro­ Larvae raised in the lab were later seen to be duced arc. Elytra: Sutural stria obsolete, occa­ cannabalistic, and Manee concluded that this sionally feebly impressed at base and apex. Disc was probably the reason for a single egg per aciculate, sparsely punctate; punctures small, chamber. Manee further noted that young larvae deep; frequently with moderately dense, weakly fed on leaf debris and later on decayed oak roots raised punctures, punctures very small to min­ and that preferred nesting sites were usually ute, very shallow, often in rows. Pygidium: Nearly near a pile of dead oak leaves blown by the wind flat in lateral view. Disc usually very finely granu­ into a hollow. late, sparsely punctate in majors, moderately Ritcher (1966) observed S. antaeus burrows in punctate in minors; punctures large, shallow, a burned-over pine woodland near Newton some with short setae in unworn specimens. Grove, North Carolina. "Each was beneath a Apical margin similarly punctate but a little more conspicuous sandy pushup. Burrows usually ex­ densely so. Genitalia: Figs. 120-121. tended obliquely, were from 8-19 inches in depth, and were provisioned with surface litter. Female.-Length 22.0-35.4 mm; width across One burrow was straight, one was L-shaped, and humerus 13.1-18.1 mm. As male except in the the rest had a single branch or were forked. Usu­ following respects: Head: Front coarsely rugose. ally a single egg was found toward the end of Clypeus with apex broadly to subacutely each burrow, or each branch of burrows .... " rounded, occasionally narrowly truncate, lacking Manee found eggs in burrows in early August, a subapical tooth, weakly reflexed; surface and Ritcher found eggs in August and Sep- A REVISION OF THE GENUS Strategus I 117 tember. Ritcher also found first instar larvae in data; Tallapoosa (1): Alexander City. ARKANAS October, second instar larvae in January, and (10): Garland (1): Hot Springs; Pulaski (6): Camp mentioned a third instar taken in the winter or Robinson, Little Rock; No data (3). CONNECTICUT spring. The composite life cycle that can be de­ (1): New Haven (1): Woodbridge. DELAWARE (1): rived (at least in North Carolina) is that eggs are Sussex (1): Dewey. FLORIDA (368): Alachua (88): deposited from early August to early September, Archer, Gainesville, High Springs; Bay (2): larvae overwinter and pupate in the spring, and Panama Beach City; Bradford (2): Starke; Bre­ adults emerge from late May to late August; adult vard (15): Cape Canaveral, Eau Gallie, Mel­ activity in North Carolina is corroborated by label bourne; Broward (5): Ft. Lauderdale; Charlotte data. (1): Punta Gorda; Clay (1): Keystone Heights; Individuals from Massachusetts and New Jer­ Collier (1): no data; Dade (73): Largo, Miami; De sey(Casey's S. septentrionis) are, with rare ex­ Soto (1): Avon Park; Duval (2): Jacksonville; ceptions, consistently very small and light red­ Hardee (1): Wauchula; Highlands (1): Archibold dish brown in color, perhaps demonstrating BioI. Stat.; Hillsborough (5): Little Manatee River, some form of ecophenotypy. In this regard, I Plant City; Indian River (1): Sebastion; Jackson suspect that further investigations into the ecol­ (1): no data; Jefferson (2): Monticello; Lake (1): ogy of the sandy pine barrens of these regions Leesburg; Lee (1): Ft. Myers; Leon (4): might yield valuable information. Harshberger Talahasee, Tall Timbers Res. Stat.; Uberty (3): (1916) has shown that the vegetation of the New Rock Bluff; Madison (2): Madison; Marion (11): Jersey pine barrens decomposes very slowly and Mcintosh, Ocala Nat'l. Forest; Okaloosa (1): may be toxic to animals because of alkaloids, Niceville; Orange (5): Oakland, Orlando, Winter­ tannins, and other phenols. Many of the plants in park; Palm Beach (7): Lake Worth; Pinellas (16): this region are poisonous, insecticidal, nervine, Cabbage Key, Clearwater, Gulfport, St. emitic, purgative, stimulatory, acrid stimulant, Petersburgh; Polk (10): Lake Alfred, Lakeland; sialagogic, diuretic, nephritic, and irritant in na­ Putname (20): Crescent City, San Mateo, Sat­ ture. Many of these compounds are commonly suma; St. Johns (4): St. Augustine; Sumpter (4): found in the soil in such areas (Burges, 1965; Inverness; Volusia (20): Brooksville, Daytona Burges et al., 1964; Whitehead, 1964), and they Beach, De Leon Springs, Enterprize, Glenwood, may definitely have a negative effect on the soil Sanford; No data (57). GEORGIA (48): Camden and litter organisms (Janzen, 1974). Feeny (1968, (1): no data; Chatham (3): Savannah; Cook (1): 1969) and Miles (1969) have demonstrated Sparks; Dekalb (27): Atlanta, Dunwoody, Ft. specific detrimental effects upon insects. McPherson; Fulton (2): Atlanta; Habersham (3): Strategus antaeus may be similarly negatively in­ no data; Hancock (1): no data; Richmond (1): fluenced under such conditions. Augusta; Thomas (3): Thomasville; Tift (1): Tif­ ton; No data (5).loWA (1): Lee (1): Keokuk. KAN­ Distribution.-Eastern United States west to SAS (3): Chautauqua (1): Sedan; Montgomery (1): Oklahoma and Texas. no data (1). KENTUCKY (3): Bell (3): Cumberland Gap; Whitley (2): Pine Knot. MARYLAND (3): Pr­ Locality Records (Fig. 4)-736 specimens ince Georges (2): Beltsville; No data (1). MAS­ examined (376 males, 360 females). Specimens SACHUSETTS (50): Barnstable (12): Cape Cod, were seen from the following collections: AMNH, Dennis, Harwhich Port, Sagamore, Truro, BCRC, CASC, CNCI, DEUN, FMNH, HAHC" INHS, Wellfleet, West Yarmouth; Dukes (3): Martha's KSUC, LACM, MCZC, OSUC, OSUO, PMNH, Vineyard; Hampden (4): Springfield; Nantucket SEMC, UC, UMMC, USNM, UWEM. (17): Nantucket; Plymouth (6): Wareham; No UNITED STATES (736).-ALABAMA (33): Dekalb data (8). MIssIssIPPI (5): Forrest (1): Brooklyn; (4): Ft. Payne, Mentone (Desoto Caves); Greene Harrison (4): Gulfport. NEW JERSEY (114): Atlantic (2): Eutaw; Jackson (2): Bryant, Pisgah; Lee (3): (4): Da Costa, Hammonton; Burlington (79): Auburn; Macon (3): Tuskegee; Madison (1): Brown's Mills, Masonville, Medford Lakes, Mt. Monte Sano; Mobile (11): Grand Bay, Mobile; Misery, Rancosas Park, Retreat; Camden (2): Montgomery (1): Montgomery; Russel (1): no Clementon; Cape May (9): Ocean City, 118 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Wildwood: Gloucester (1): Wenonah; Meddlesex totype label. Also one male paralectotype at (1): Jamesburg; Ocean (16): Lakehurst, Man­ ZMHU. Type locality: Cordoba, Argentina.] ahawkin, Seaside Park, Whiting; No data (2). NEW YORK (8): Suffolk (3): Greenport, Riverhead; Male.-Length 31.0-42.0 mm; width across No data (5). NORTH CAROLINA (27): Halifax (1): humerus 18.0-21.0 mm. Color castaneous to Halifax; Johnston (1): Benson; Moore (22): Eagle piceous, shining. Head: Front rugose to coarsely Springs, Eastwood, Pinehurst, Southern Pines: punctate; punctures large, shallow, confluent. Robeson (1): Maxton; Wake (2): Raleigh. Clypeus with apex truncate, moderately emargi­ OKLAHOMA (5): Caddo (1): Anadarko; Kingfisher nate; surface rugulose to rugose. Tubercles con­ (1): Kingfisher; Logan (1): Guthrie; Tulsa (2): ical, small to moderate in size, slightly trans­ Tulsa. PENNSYLVANIA (3): Montgomery (3): verse, usually distinctly separated. Mandibles Frankford. RHODE ISLAND (14): Kent (5): Button­ with basal lobe small, rounded; middle lobe wood,s; Warwick; Providence (9): Providence. large, triangular, apex rounded; apical lobe SOUTH CAROLINA (24): Aiken (6): Aiken; Beaufort small, triangular. Interocular width 3.0 transverse (1); Hilton Head Island; Charleston (1): Santee eye diameters. Mesosternum: Anterior half Plantation (South Santee River); Chesterfield (1): setigerously punctate. Pronotum: Base with a McBee; Colleton (1): Hendersonville; Fairfield wide rugose band; band reduced medially nearly (1): W,oodward; Florence (1): Florence; Horry (1): to basal bead. Disc aciculate, punctate; Arrowhead Lake (Myrtle Beach); Jasper (2): punctures small and minute mixed, moderate in Bluffton; Richland (3); Columbia; No data (6). density, deep. Sides similar to disc except TENNESSEE (21): Grundy (2): Camp Mt. Lake; punctures larger; lateral margin with a band of Hamilton (2): Signal Mountain; Knox (2): Knox­ moderate to large punctures, punctures moder­ ville; Lake (1): Tiptonville; Madison (1): no data; ately dense, simple to crescent-shaped, shallow. Morgan (12): Burrville, Deer Lodge, Sunbright; Anterior angles and fovea either side of middle No data (1). TEXAS (14): Dallas (3): Dallas; Leon as disc to rugose. Fovea deep, with or without a (3): no data; Nueces (2): Corpus Christi; No data low, feeble, median, longitudinal ridge extending (6). VIRGINA (4): James City (1): Newport News; posteriorly from base of anterior horn. Horns: Nelson (1): no data; Southampton (1): Franklin; Anterior short to moderate, stout, attenuate, No data (1). February (2), March (4), April (16), curving forward and upward; apex narrowly May (35), June (143), July (198), August (118), rounded, occasionally very feebly emarginate; September (44), October (13), November (2), De­ dorsal surface rounded to nearly flat. Posterior cember (4). horns vary from low, rounded bosses to very short, triangular, stout, laterally compressed Remarks.-The absence of a sutural stria will horns; in dorsal view horns parallel, bases joined easily separate S. antaeus from all other species across disc in a shallow arc. Elytra: Sutural stria in the genus. Strategus antaeus is somewhat var­ strongly impressed, weakly crenulate. Disc fre­ iable with regard to shape, size, and color. Spec­ quently wrinkled, aciculate, punctate; punctures imens from the southern part of the range are minute, small and moderate mixed, moderately generally darker and larger than those from the dense, shallow, occasionally in several incom­ northern part of the range. plete rows; lateral half of disc frequently with 3-5 Strategus antaeus houstonensis Knaus (1925) shallow, incomplete striae; striae occasionally is not a valid subspecies as proposed but merely produced into broad, shallow furrows; furrows one aspect of the intraspecific variation seen interrupted, apically branched. Sides similar to within the species. disc except punctures slightly larger, striae ab­ Strategus argentinus Kolbe sent, and wrinkling usually pronounced. Apex (Figs. 8, 32-33,122-123) densely punctate, punctures as on disc. Pygidium: Convex in lateral view. Disc aciculate, Strategus argentinus Kolbe, 1906: 24. [Lectotype usually finely roughened to very finely granulate, male, labeled "Argentinien, Cordoba, J. Fren­ punctate; punctures small, moderately dense, zel's," here designated; at ZMHU with my lec- very shallow; those on apical midline large, A REVISION OF THE GENUS Strategus / 119 sparse, deep, setigerous in unworn specimens; tinguish it from all other species. From the spec­ punctures often greatly reduced. Apical margins imens examined, it apparently does not possess densely punctate to rugose. Genitalia: Figs. what could be called a major stage of develop­ 122-123. ment with regard to armature. S. argentinus is an uncommon species. Female.-Length 30.5-35.0 mm; width across humerus 14.7-17.0 mm. As male except in the following respects: Head: Clypeal apex nearly Strategus at/anticus, new species rectangular in unworn specimens, moderately (Figs. 1, 34-35, 124-125) reflexed; apex narrowly to broadly rounded or truncate in worn specimens. Tubercles trans­ Type Material.-Holotype male, labeled "WaIt­ verse, usually connected by a weak carina. Man­ ing (sic) Island, Bahamas, Oct. 10-21. 91, C. B. dibles similar to those of male but smaller, espe­ Cory (?)," deposited at MCZC. cially middle lobe. Pronotum: Base with rugose Holotype.-(Figs. 34-35, 124-125).-Male. band Rarrow to moderate. Sides with punctures Length 32.0 mm; width across humerus 15.4 mm. moderately large, dense, grading into rugosity Color dark castaneous, feebly shining. Head: anteriorly at about middle; a slightly depressed, Front distinctly concave, basal half finely granu­ rounded area of dense punctures postero-Iateral late, apical half densely punctate; punctures of fovea. Anterior third rugose. Fovea shallow. large, shallow, some confluent; a trace of setae Tubercle strong, conical, transverse. Pygidium: above eye. Clypeus with apex broadly rounded, In lateral view basal half convex, apical half feebly reflexed; surface finely granulate, densely nearly flat. Disc punctate; punctures small and punctate; punctures large and minute mixed, large mixed, moderately dense, shallow; large shallow. Tubercles conical, transverse, very low, punctures simple to oblong, setigerous in un­ widely separated. Mandibles with basal lobe worn specimens. Lateral emargination shallow, small, rounded; middle lobe subtriangular, more strongly punctured. small, apex nearly right angled; apical lobe Biology.-Unknown. small, obtusely rounded. Interocular width 2.33 transverse eye diameters. Mesosternum: Com­ Distribution.-Uruguay and northeast Argen­ pletely and setigerously punctate. Pronotum: tina. Endrodi (1973b) reported 4 specimens from Base with a few moderate, elongate punctures Bolivia, but without seeing the specimens I am either side of midline. Disc extremely finely sub­ doubtful about this record. granulate, sparsely punctate; punctures small, Locality Records (Fig. 8).-23 specimens moderately deep. Sides in basal 21.3 similar except examined (16 males, 7 females). Specimens were punctures a little larger and a little more dense, seen from the following collections: BMNH, especially near lateral margin. Anterior third FMNH, MCZC, UC, URU, USNM, USP, ZMHU. rugo-punctate. Fovea virtually effaced, only an extremely shallow, barely noticeable depression ARGENTINA (16).-CATAMARCA (2): Catamarca; remains. Horns: None; even anterior tubercle C6RDOBA (4): Cordoba; SALTA (2): Coronel absent. Elytra: sutural stria strongly impressed, Moldes, Gran Chaco; SANTIAGO DEL ESTERO (4): crenulate. Disc finely granulate, punctate; Banda, Chalya; No data (4). February (2). punctures small and large mixed, small November (5), December (1). ' punctures sparse (becoming more so laterally), URUGUARY (7).-ARTIGAS (2): Capital, Catalan shallow; large punctures ocellate-umbilicate, Chico; COLONIA (1): Piedra de los Indios, moderately dense, shallow, and irregular except PAYSANDU (1): Rt. 3 (Km. 383); SALTO (2): San for a regular row in center of disc and a regular Antonio; TACUAREMB6 (1): Puntas Arroyo double row on lateral margin of disc. Sides simi­ Lau reles. January (1), December (6). lar except wrinkled behind humerus, small punctures very sparse, large punctures sparse, a Remarks.-Although S. argentinus is a rather double row of moderate punctures in middle. nondescript species, the key characters will dis- Apex moderately punctate; punctures moderate 120 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

in size, shallow. Pygidium: Very strongly convex shining. Head: Front largely hidden by closely in lateral view, protuberant at middle. Surface appressed anterior pronotal horn; surface finely finely granulate, sparsely punctate, punctures granulate, feebly rugo-punctate (especially be­ small, shallow. Genitalia: Figs. 124-125. hind tubercles). Clypeus with apex rounded and weakly emarginate, moderately reflexed; surface Female.-Unknown. finely granulate, feebly rugo-punctate. Tubercles conical, nearly effaced. Mandibles with basal Biology.-Unknown. lobe submoderate, prominently rounded; middle lobe moderate in size, subtriangular, apex Distribution.-Watling Island (San Salvador), rounded; apical lobe subtriangular, slightly smal­ Bahamas. ler than middle lobe, apex acute. Interocular .Locality Records (Fig. 1 ).-One specimen width 1.66 transverse eye diameters. Mesoster­ .examin~d (holotype). Specimen deposited at num: Anterior half setigerously punctate. Pro­ MCZC. notum: Base with a narrow, rugose band; band reduced medially almost to basal bead. Disc very BAHAMAS (1 ).-WATLING ISLAND. October (1). finely granulate, punctate; punctures moderate and minute mixed; moderate punctures moder­ ,. Remarks.-The completely punctate mesos­ ately dense, deep (becoming shallower laterally) ternum and the presence of distinct rows of minute punctures sparse, shallow. Sides in basal punctures on the elytral disc will distinguish this half with punctures a little less dense, slightly species from al\ others. It is unique among larger to large, shallow; sides in anterior half and Strategus in that the pronotal fovea and tubercle anterior angles sparsely punctate, punctures are virtually obsolete. The other characters and small, shallow. Fovea extremely shallow, the genitalia in particular place it in this genus. sparsely punctate, punctures small to very small, Zoogeographically, it is the most isolated of the shallow. Horns: Anterior long, robust, dorso­ species and is probably derived from close rela­ ventrally compressed, expanding apically to a tives in Cuba or Hispaniola such as the ancestors very strongly forked apex, each prong of fork of S. aenobarbus, S. ajax, or S. talpa. acutely rounded and divided by a deep U-shaped Etymology.-From the Latin atlas, a mythical emargination; in lateral view horn curves for­ god who held up the heavens; here named for its ward and downward closely over head and then occurrence on San Salvador in the Atlantic bends upward at about middle; dorsal surface ocean. flat to slightly concave, each lateral margin lon­ gitudinally carinate. Posterior horns completely lacking. Elytra: Sutural stria impressed, crenu­ late. Disc wrinkled, finely granulate, punctate; Strategus cayman;, new species punctures small and minute mixed; small (Figs. 1, 36-39, 126-127) punctures moderate in density, shallow, and be­ Type material.-Holotype male, labeled "29 v. coming increasingly ocellate-umbilicate laterally 1938, Little Cayman, S. Coast of South Town and basally; minute punctures dense, very shal­ light trap A," "17. iv-26. viii 1938, Oxf. Un: low; 3 shallow, incomplete striae on lateral half Cayman Is. BioI. Exped., Coil. by C. B: Lewis, G. of disc. Sides similar to disc except without H. Thompson," "Strategus simson (L.), local striae, with increased wrinkling behind humerus, form. det. 1950, E. A. Chapin." Allotype female and ocellate-umbilicate punctures moderate to with same data as holotype. Types deposited at large. Apex densely punctate; punctures small to BMNH; 18 males and one female paratypes de­ moderate, simple to ocellate-umbilicate, shal­ posited in BCRC, BMNH HAHC SEBO EN- low. Pygidium: Convex in lateral view. Disc very DROOL " finely subgranulate, sparsely punctate; punctures small to moderate, very shallow. Api­ Holotype.-Male. Length 38.1 mm; width cal margins rugulose either side of midline. across humerus 18.0 mm. Color piceous, feebly Genitalia: Figs. 126-127. A REVISION OF THE GENUS Strategus / 121

Allotype.-Female. Length 37.0 mm; width ac­ Locality Records (Fig. 1 ).-21 specimens ross humerus 17.7 mm. Color slightly lighter examined (19 males, 2 females). Specimens de­ than holotype. As holotype except in the follow­ posited in the following collections: BCRC, ing respects: Head: Front finely granulate, BMNH, HAHC, SEBO ENDRODI. strongly rugo-punctate to rugose. Clypeus with CAYMAN ISLANDS (21 ).-CAYMAN BRAe (6): apex broadly rounded, very weakly emarginate; north coast Stakes Bay, west end of Cotton-tree surface rugose. Mandibles similar to those of land; LITTLE CAYMAN (15): south coast of South holotype but slightly smaller. Pronotum: Base Town. May (20), June (1). with a narrow rugo-punctate band, band re­ duced nearly to basal bead at midline; basal Remarks.-The partially punctate mesoster­ fourth of midline with an irregular, longitudinal num, rounded and weakly emarginate clypeal patoh of moderate to large, shallow punctures. apex, dull elytra, distinctly forked anterior horn, Sides in basal half moderately densely punctate; lack of posterior horns, and an interocular width punctures moderate to large, shallow; lateral of less than two transverse eye diameters will margin in basal half with a narrow rugose band; serve to separate S. caymani from any other just postero-Iateral of fovea is a slightly depres­ species. In 1950 Chapin concluded that the type sed, rounded patch of rugosity. Anterior half series in the BMNH was a local form of S. sim­ rugo-punctate to rugose. Fovea nearly obsolete, son. Even though the genitalia are almost identi­ faintly carinate longitudinally down middle. cal, I believe other characters are sufficiently dis­ Tubercle conical, transverse, nearly effaced. tinctive to justify new species status. S. caymani Pygidium: Strongly convex and protuberant at is very closely related to S. simson. middle in lateral view. Disc very finely subgranu­ Etymology.-This species is named after the late, moderately densely punctate, punctures Cayman Islands on which it occurs. small and minute mixed, very shallow. Lateral emargination shallow, rugulose. Variation.-Males (18 paratypes): Length 28.7-39.4 mm; width across humerus 13.4-18.0 Strategus centaurus Kolbe mm. Head: Interocular width 1.6&-2.0 transverse (Figs. 9, 40-43, 128-129) eye diameters. Mesosternum: Anterior 1/2-% setigerously punctate. Pronotum: As holotype to Strategus centaurus Kolbe, 1906: 29. [Types not base either side of middle with large, simple to seen. Holotype and presumably allotype at oblong punctures. Anterior half with punctures ZMHU. Type locality: Brazil.] moderate in size. Horns: Majors (Figs. 3&-37) with anterior narrowly to very widely forked at Male.-Length 50.5-80.0 mm; width across apex, horn subslender to very stout. Minors humerus 24.8-37.3 mm. Color piceous to black, (Figs. 38-39) with anterior short, expanded very shining. Head: Front coarsely rugo-punctate to little apically, apex triangularly notched; dorsal rugose. Clypeus with apex strongly reflexed, very surface rounded to weakly carinate longitudi­ broadly truncate, and with a moderate to very nally down middle or down middle and on lateral deep, median, triangular excision; surface usu­ margins. ally rugose in minors, moderately punctate in Females (1 paratype): Length 33.2 mm,; width majors; punctures large, shallow. Tubercles con­ across humerus 15.3 mm. Head: Interocular ical, strong, transverse, distinctly separated to width 2.0 transverse eye diameters. Pronotum: weakly connected in minors by a weak, trans­ Fovea obsolete. verse carina. Mandibles with basal lobe small, prominently rounded; middle lobe very large, Biology.-Label data indicate "attacked by a triangular, apex rounded; apical lobe small, tachinid" and "taken at lights"; otherwise un­ triangular, apex usually acute. Interocular width known. 2.0-2.5 transverse eye diameters. Mesosternum: Distribution.-Little Cayman Island and Anterior % setigerously punctate. Pronotum: Cayman Brac (West Indies). Base with transverse, rugose band wide to very 122 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

wide; band reduced medially almost to basal Female.-Length 52.5-64.9 mm; width across bead. Disc aciculate, sparsely punctate; humerus 25.6-31.5 mm. As male except in the punctures small, deep. Sides of minors similar to following respects: Head: Clypeus with apex disc except lateral margin rugo-punctate in basal rounded to broadly truncate, moderately re­ half, grading to rugose in anterior half; a strong flexed. Tubercles transverse, usually connected rugo-punctate patch present at base of posterior by a weak, transverse carina. Mandibles similar horns or bosses; sides of majors also similar to to those of male but smaller, especially middle disc except marginal band reduced to a few lobe. Interocular width 2.33-3.0 transverse eye sparse, moderately large punctures, and a diameters. Pronotum: Sides in basal half rugo­ rugo-punctate patch present at base of posterior punctate to densely punctate; punctures moder­ horns. Anterior half rugose except in very large ate to large, deep, frequently confluent; a slightly males, and then rugose only in anterior angles. depressed, rounded patch of rugosity present H<;>rns: Majors (Figs. 40-41) with anterior long, just postero-Iateral of fovea. Anterior half stout, attenuate, curving forward and upward, rugose. Fovea moderately deep. Tubercle coni­ , apex narrowly rounded, dorsal surface rounded. cal, strong, transverse. Pygidium: Occasionally Posterior horns long, stout, not attenuate, later­ as in male, more commonly entirely rugose; ally compressed, projecting forward and upward sparse setae present apically in unworn speci­ at':about 50-80° from plane of disc; apex ob­ mens. liquely truncate, occasionally weakly to moder­ ately scalloped and with posterior flaring in Biology.-Unknown. Costa Lima (1953) re­ well-developed specimens; in dorsal view horns ported that they were a pest of palms at Bahia, subparallel to curving toward one another. Brazil. Minors (Figs. 42-43) with anterior as in majors except short. Posterior horns vary from low, Distribution.-South America. The single rec­ subparallel to slightly divergent pyramidal bos­ ord from Venezuela (USNM) is somewhat doubt­ ses to short, stout horns; horns attenuate, later­ ful. ally compressed, apex broadly rounded, project­ ing vertically to subvertically from plane of disc. Locality Records (Fig. 9).-28 specimens Elytra: Sutural stria strongly impressed, crenu­ examined (13 males, 15 females). SpeCimens late. Disc aciculate, sparsely punctate; were seen from the following collections: AMNH, punctures small and minute mixed; median Y3-Y2 BMNH, CNCI, FMNH, HAHC, MCZC, UC, USNM. frequently with sparse, ocellate punctures; ARGENTINA (2).-MISIONES (2): EI Dorado, punctures small to moderate; in addition lateral Puerto Bemberg. November (2) Y~h usually with 6 rows of moderate to large, ocellate punctures, rows 1-2 and 4-5 regular and BRAZIL (32).-BAHIA (7): Bahia, 20 mi. SW. Sal­ distinct, rows 3 and 6 irregular and confused; vador, no data; ESPIR'iTO SANTO (2): Espirito rows 2, 4, and 5 frequently in or adjacent to a Santo; PARA (4): Para, Santarem; PARANA (3): shallow, incomplete stria; rows of punctures and Caviuna, Villa Velha; SANTA CATARINA (6): size of punctures occasionally reduced, espe­ Corupa, Nova Teutonia, no data; SAo PAULO (1): cially row 1. Sides with 3-6 confused rows of Piracicaba; No data (9). January (2), April (3), punctures; punctures ocellate, moderate to June (1), November (1), December (3). large, becoming obsolete posteriorly. Apex PARAGUAY (3).-lnsufficient data (3). January sparsely to moderately punctate; punctures (2), December (1). small, shallow. Pygidium: In lateral view basal half convex, apical half weakly concave. Surface Remarks.-The large mandibles, the definite aciculate, varies from weakly rugo-punctate to rows of moderate to large ocellate punctures on punctate; punctures small to moderately large, the lateral half 9f the elytral disc, the strongly simple to oblong, shallow, sparse, becoming excised clypeal apex, and a length greater than denser near apical margins either side of middle. 50 mm will serve to separate this, the largest of Genitalia: Figs. 128-129. all Strategus, from any other species. A REVISION OF THE GENUS Strategus / 123

Strategus cessatus Wickham Anastrategus durangoensis Casey, 1915: 234. (Figs. 6, 174) [Holotype male at USNM. Type locality: Tepehuanes, Durango, Mexico.] Strategus cessatus Wickham, 1914: 461. Anastrategus inflatus Casey, 1915: 234. [Holotype at MCZC. Type locality: Florissant, [Holotype male at USNM. Type locality: Colorado, United States.] Arizona, United States.] Anastrategus tantalus Casey, 1915: 235. [Lec­ totype male, labeled "Oslar, Prescott, Ariz.," Remarks.-The reader is referred to Wickham "Casey bequest 1923," Type USNM 48627," for a description of this, the only known fossil and "tantalus Csy.," here designated; at USNM Strategus. The single elytron was originally de­ (Casey collection) with my lectotype label. Lec­ scribed as being Miocene, but subsequent toallotype female, labeled as male except for studies have established that the Florissant de­ type label which reads "tantalus - 3, Paratype posits are actually Oligocene. My examination of USNM 48627," here designated; at USNM with the type initially left some doubt as to whether my lectoallotype label. Also one male paralec­ the specimen was indeed a Strategus, especially totype at USNM. Type locality: Prescott, since there was only one elytron to work with. Arizona, United States.] Wickham placed it in Strategus because he thought that it corresponded more to this genus Male.-Length 24.5-41.0 mm; width across than to any other related genera, and in this re­ humerus 11.0-20.0 mm. Color castaneous to gard I now agree with Wickham. black, shining. Head: Front rugose to very Strategus cessatus is closely related to S. ces­ coarsely punctate; punctures dense, large, deep, sus and probably occupied a habitat comparable often confluent; setigerous above eye in unworn to today's S. cessus. specimens. Clypeus with apex narrowly rounded (rarely feebly emarginate), moderately reflexed. Tubercles strong, conical, transverse, joined by a Strategus cessus LeConte moderate to strong, transverse carina. Mentum (Figs. 6, 44-45, 130-131) with disc coarsely punctate; punctures large, dense, often confluent. Mandibles nearly square Strategus cessus LeConte, 1866: 382. [Lectotype in dorsal view. Interocular width 4.0-4.5 trans­ female, labeled "Ariz.," "S. cess us LeC., verse eye diameters. Mesosternum: Anterior half Coves/3," and "Type/3780," here designated; setigerously punctate. Pronotum: Base with a at MCZC with my lectotype label. Lectoallotype narrow, coarsely rugo-punctate band. Disc male, labeled "Ariz.," here designated; at aciculate, punctate; punctures moderately dense ICCM (Ulke Collection) with my lectoallotype (sparser along midline), moderate to large, shal­ label. Type locality: Arizona, United States.] low to deep; usually a feeble, longitudinal Strategus beckeri Kolbe, 1906: 14. [Lectotype depression along midline. Sides with large, de­ male, labeled "Mexico, J. Flohr G.," "70808," nse, moderately deep punctures, becoming and "beckeri n. sp.," and lectoallotype female, coarsely rugo-punctate to rugose in anterior labeled as lectotype but without number, here quarter. Anterior third rugose. Fovea moderately designated; at ZMHU with my lectotype labels. deep. Horns: Anterior a moderate to strong Also one male paralectotype at ZMHU. Type tubercle; tubercle conical, very transverse, apex locality: Sierra Madre, Durango, Mexico. Lec­ usually distinctly emarginate, truncate in worn totype illustrated in Kolbe.] New synonymy. specimens. Posterior horns completely lacking. Anastrategus cess us cavicauda Casey, 1915: Elytra: Sutural stria weakly to moderately 233. [Lectotype female here designated; at impressed, feebly crenulate; anterior fifth occa­ USNM with my lectotype label. Also one female sionally effaced and replaced by moderate-sized, paralectotype at USNM. Type locality: un­ shallow, irregularly shaped punctures. Disc known, but Casey suspected New Mexico, aciculate, punctate; punctures sparse to moder­ United States.] ately dense, small to large, shallow; 2-3 124 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM punctures frequently joined to form an irregular, San Pedro de Guanacevi, Sierra Madre; JALISCO transverse furrow; lateral half of disc with 2 dou­ (1): no data; No data (2). June (1), July (1), August ble rows of impressed, occasionally interrupted, (11 ). punctate striae; punctures moderate to large, UNITED STATES (154).-ARIZONA (138): Apache deep; striae variably effaced, especially the first. (2): Springville; Cochise (59): Carr Canyon (15 Sides as disc except usually with 1 vague, double mi. S. Sierra Vista), Cave Creek Ranch, Cochise row of feebly impressed, punctate striae adja­ Stronghold, Huachuca Mts., Miller Canyon cent to disc; punctures moderate in size; striae (Huachuca Mts.), Paradise, Parker Canyon Lake, often reduced to completely absent; area just Portal, Southwest Research Station (Portal), behind humerus transversely wrinkled or with a Sunnyside; Gila (7): base of Pinal Mts., Globe, weak to strong row of moderate to large, deep Payson; Maricopa (4): Phoenix: Navaho (2): punctures. Apex punctate; punctures dense to Kayenta, Snowflake; Pima (5): Madera Canyon, rugo-pun9tate. Pygidium: In lateral view basal Santa Rita Mts.; Santa Cruz (34): Gardner Can­ third stro-ngly convex, apical % nearly flat. Disc yon, Hidden Springs Valley (9 mi. E. Sanoita), very finely granulate, aciculate, punctate; Madera Canyon, Nogales, 15 mi. NW. Nogales, punctures sparse to moderately dense and Peiia Blanca Lake, Sondida Creek (9 mi. S. coarse, small to large, shallow; a few short setae Patagonia), Tubac, west slope Patagonia Mts.; near apex in unworn specimens. Genitalia: Figs. Yavapai (18): Congress, Prescott; No data (7). 130-131:' NEW MEXICO (15): Bernalillo (1): Albuquerque; - Female.-Length 24.8-37.2 mm; width across Catron (2): Apache Nat'l. Forest, Reserve; Grant humerus 10.0-19.0 mm. As male except in the (5): Ft. Bayard, Silver City; San Miguel (2): La following respects: Pygidium: In lateral view Trementina (Fall and Cockerell, 1907), Las basal half convex, apical half concave. Disc Vegas; No data (5). April (1), May (2), June (16), aciculate or not, sparsely punctate; punctures July (86), August (41), September (6). shallow, usually small, occasionally several Remarks.-The nearly square mandibles are moderately large. Apical margin densely diagnostic for S. cessus. Because the males lack punctate; a few short setae present in unworn horns, the males and females more closely re­ specimens; lateral emargination absent. semble each other in this species than in any Biology.-Unknown. Label data indicate that other. In the presence of larger samples it is now S. cessus has been taken at lights and from apparent that the diagnostic characters given by 1300-2736 meters in altitude. Kolbe (1906) for S. beckeri are merely part of the normal variation found in S. cessus. It is interest­ Distribution.-Southwest United States and ing to note, however, that virtually all of the spec­ northwest Mexico. The single record from imens from Durango in Mexico are considerably California is possibly erroneous or due to con­ lighter in color than those from the rest of the veyance by man. range. Locality Records (Fig. 6).-184 specimens examined (66 males, 110 females). Specimens Strategus craigi, new species were seen from the following collections: AMNH, (Figs. 6, 46-47, 132-133) BCRC, BMNH, CASC, CNCI, DEUN, FMNH, HAHC, ICCM, KSUC, LACM, MCZC, PMNH, Type Material.-Holotype male, labeled OSUC, OSUO, SEMC, UMMC, USNM, ZMHU. "Jalapa, Mexico, M. Trujillo," "B.C.A., Col., 11(2), Strategus julianus," and "~"; depOSited at MEXICO (29).-CHIHUAHUA (16): 5 mi. N. Cerro BMNH. Campana (Sierro del Nido), 8 mi. W. Matachic, 10 mi. W. Namiquipa, 15 mi. E. Parral, Santa Bar­ Holotype (Figs. 46-47).-Male. Length 37.7 bara, Santa Clara Canyon (5 mi. W. Barrita); mm; width across humerlls 18.6 mm. Color pice­ DURANGO (10): 3 mi. W. Durango, 25 mi. W. ous, feebly shining. Head: Front grossly rugo­ Durango, Javilanes, Otinapa, Palos Colorados, punctate, no supra-ocular setae seen. Clypeus A REVISION OF THE GENUS Strategus / 125 with apex broadly rounded, subtruncate, moder­ Etymology.-This species is named in honor ately reflexed; surface coarsely rugo-punctate. of my younger brother. Tubercles conical, transverse, connected by a strong, transverse carina. Mentum with sparse, small to moderate punctures on disc. Mandibles Strategus fallaciosus Kolbe with basal lobe moderate in size, prominently (Figs. 6,48-49,134-135) rounded; middle lobe moderate in size, subequal with basal lobe, also rounded; apical lobe small, Strategus fallaciosus Kolbe, 1906: 16. [Lectotype rounded. Interocular width 4.0 transverse eye male, labeled "Mexico, Motzerongo (sic), Staat diameters. Mesosternum: Anterior half setiger­ Veracruz, R. Becker," "70832," "Type," and ously punctate. Pronotum: Base with a wide, "fallaciosus," here designated; at ZMHU with rugo-punctate to punctate band; punctures my lectotype label. Also one male paralec­ large, dense, shallow, reduced almost to basal totype at ZMHU. Type locality: Motzorongo, bead at midline. Disc aciculate, punctate, Veracruz, Mexico. Lectotype illustrated in punctures moderately dense, small, shallow; a Kolbe.] V-shaped band of moderate to large, shallow punctures extending from base and forking at Male.-Length 43.5-45.0 mm; width across base of fovea, arms extending obliquely forward; humerus 20.8-22.5 mm. Color dark castaneous, a 'slightly depressed, rounded area of rugosity feebly shining. Head: Front grossly rugo­ just postero-Iateral of fovea. Remainder of pro­ punctate, setigerous above eye in unworn spec­ notum rugose. Fovea deep. Horns: Anterior a low imens. Clypeus with apex rounded in worn spec­ tubercle, tubercle conical, transverse, apex sub­ imens, extended as a rounded to squared tooth truncate. Posterior horns lacking. Elytra: Sutural in unworn specimens, slightly reflexed; surface stria strongly impressed, crenulate. Disc acicu­ rugo-punctate to rugulose. Tubercles strong, late, weakly wrinkled, mode~ately punctate; conical, transverse, connected by a strong, punctures small and minute mixed, shallow; 3 transverse carina. Mandibles with basal lobe feebly impressed, incomplete striae on lateral 21.3. small, prominently rounded; middle lobe large, Sides similar except sparsely punctate. Apex triangular, apex very acute; apical lobe similar to densely punctate; punctures small to moderate, middle lobe but smaller. Interocular width 3.33 shallow. Pygidium: In lateral view basal half con­ transverse eye diameters. Mesosternum: An­ vex, apical half nearly flat. Disc roughened, terior half setigerously punctate. Pronotum: moderately punctate; punctures minute, very Base with a wide, rugo-punctate to rugose band; shallow, becoming denser and larger in area of band reduced nearly to basal bead at midline. lateral emargination; a few sparse, large Disc faintly impressed medially, aciculate, punctures near midline; a few short setae near punctate; punctures moderately dense, small to apex. Genitalia: Figs 132-133. moderate in size, becoming larger at base of fovea, deep. Sides in basal half similar to disc Biology.-Unknown. except punctures moderately large and lateral margin with a narrow rugo-punctate to rugose Distribution.-Mexico. band; sides grade to rugo-punctate and then to rugose in anterior half; a slightly depressed, Locality Records (Fig. 6).-One male rounded area of rugosity postero-Iateral of fovea. examined (holotype). Deposited at BMNH. Anterior third rugose. Fovea subquadrate, deep. MEXICO (1).-VERACRUZ (1): Jalapa. Horns: Anterior a large, erect tubercle; tubercle conical, transverse, apex very feebly emarginate. Remarks.-This species is very closely related Posterior horns are very low, rounded bosses; in to S. cessus but may be distinguished from it by dorsal view bases joined across disc in a shallow the less punctate mentum, distinctly lobed man­ arc; arc strongly' produced at middle. Elytra: dibles, and a distinct, wide, rugo-punctate band Sutural stria usually moderately impressed and at the base of the pronotum. crenulate, occasionally reduced to a row of ir- 126 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

regularly shaped, moderate to large, deep Strategus fascinus Burmeister punctures. Disc aciculate, punctate; punctures (Figs. 8, 50-52, 136-137) sparse to moderate, small, deep; usually 2-3 very feebly impressed, incomplete striae on lateral Strategus fascinus Burmeister, 1847: 131. half. Sides wrinkled behind humerus, sparsely to [Holotype male at Martin Luther Universitat, moderately punctate;. punctures small to moder­ Halle, German Democratic Republic. Type loc­ ate, shallow. Apex densely punctate to rugo­ ality: Colombia.] punctate. Pygidium: Convex in lateral view. Disc finely granulate, very densely punctate; Male.-Length 33.4-38.4 mm; width across punctures small to moderately large, shallow; humerus 16.4-18.7 mm. Color castaneous to usually a few large, shallow punctures on mid­ piceous, shining. Head: Front weakly (but dis­ Ii~Et; apical margin with small, sparse setae in tinctly) to strongly rugo-punctate, setigerous unworn specimens. Genitalia: Figs. 134-135. above eye. Clyeus with apex weakly to moder­ ately excised, moderately reflexed; surface Female.-Length 40.0-43.8 mm; width across rugo-punctate to sparsely punctate; punctures humerus 18.7-22.0 mm. As male except in the small to moderate, shallow. Tubercles strong, following respects: Head: Apex of clypeus as conical, slightly transverse, widely separated. male to extended as an acute tooth. Mandibles Mandibles with basal lobe small, prominently with middle lobe slightly smaller than in male. rounded; middle lobe moderately large, triangu­ Pronotum: Discal punctures usually do not be­ lar, apex rounded; apical lobe small, triangular. come larger at base of fovea. Fovea rounded. Interocular width 2.33-2.66 transverse eye Tubercle conical, low, transverse; apex weakly diameters. Mesosternum: Completely and but distinctly emarginate. Pygidium: In lateral setigerously punctate, often weakly so in poste­ view basal half convex, apical half concave. Disc rior half. Pronotum: Base with a narrow rugose finely and entirely setigerous in unworn speci­ band; band reduced nearly to basal bead at mid­ mens. Apical margins densely punctate to rugo­ dle. Disc aciculate, punctate; punctures very punctate; lateral emargination extremely shal­ sparse, small, shallow. Sides as disc except mar­ low. gin in basal half usually rugo-punctate to Biology.-Unknown. Label data indicate spec­ punctate in a narrow band; punctures small to imens have been found dead in a sawdust pile moderately large; less-developed individuals with a patch of moderate-sized, shallow and at elevations of 1,335 and 1,435 m. punctures at base of posterior horns. Anterior Distribution.-Mexico. half as disc. Fovea deep, median longitudinal ridge from base of anterior horn very low and Locality Records (Fig. 6).-7 specimens broadly rounded, not prominent. Horns: Majors examined (3 males, 4 females). Specimens were (Figs. 50-51) with anterior long, slender, sides seen from the following collections: CNCI, subparallel (not attenuate), curving forward and SEMC, ZMHU. upward, apex slightly expanded and weakly to MEXICO (7).-JALISCO (2): 3 mi. SW. Mazamitla, moderately triangularly notched; dorsal surface 12.4 mi. S. Tecalitlan; SONORA (3): Yecora; VER­ round to flat, usually a small, longitudinal, me­ ACRUZ (2): Motzorongo. June (1), August (1), dian carina on apical half in large specimens. November (3). Posterior horns long, slender, attenuate, laterally compressed, extending forward and upward at Remarks.-The acutely triangular second and about 20-40° from plane of disc; apex acutely third lobes of the mandibles together with the rounded; occasionally a Slight enlargement on large, elongate, subparallel body shape will usu­ ventral edge about half the distance from apex; ally serve to distinguish this species at a glance. in dorsal view horns subparallel to slightly di­ This species is apparently very rare. The female verging to slightly curved toward one another; is recognized and described here for the first bases not appreciably joined across disc in an time. arc except in smaller specimens. Minors (Fig. A REVISION OF THE GENUS Strategus / 127

52): True minors not seen. Intermediate speci­ Curran"; deposited at USNM. Allotype female, men illustrated similar to majors except horns labeled "Hamburg Farm, Reventazon, Limon, reduced. Very small individuals presumably with Costa Rica, May 3, 1928, F. Nevermann"; depos­ typical reduction in armature and increase in ited at USNM. Also four male and 11 female sculpturing. Elytra: Sutrual stria impressed, cre­ paratypes deposited in the following collections: nulate. Disc aciculate, sparsely punctate; AHCC, AMNH, BCRC, BMNH, DEUN, FMNH, punctures small, deep. Sides similar except with HAHC. 2 short rows of ocellate-umbilicate punctures behind humerus; punctures moderately large, Holotype (Figs. 53-54).-Male. Length 52.7 shallow, rows occasionally reduced. Apex den­ mm; width across humerus 25.5 mm. Color pice­ sely punctate; punctures sparse to moderate in ous, shining. Head: Front rugose, setae above density, small, shallow. Pygidium: Convex in lat­ eye absent (probably worn off). Clypeus with eral view. Disc finely subgranulate, virtually im­ apex broadly truncate, shallowly emarginate, punctate to very sparsely punctate; punctures moderately reflexed; surface aciculate, moder­ small, shallow, more distinct on apical margins ately punctate; punctures small, very shallow. and midline of disc. Anterior margins occasion­ Tubercles conical, strong, widely separated. ally with a narrow band of rugosity either side of Mandibles with basal lobe small, very promi­ mipdle. Genitalia: Figs. 13&-137. nently rounded; middle lobe large, triangular, apex rounded; apical lobe similar to basal lobe. Female.-Apparently unknown. Interocular width 2.5 transverse eye diameters. Biology.-U nknown. Mesosternum: Anterior half setigerously punctate. Pronotum: Base with a wide rugose Distribution.-Colom bia. band; band reduced almost to basal bead at Locality Records (Fig. 8).-9 specimens midd Ie. Disc aciculate, sparsely punctate; examined (all males.) Specimens were seen from punctures small, shallow. Sides as disc except a the following collections: AMNH, BMNH, UNC, small patch of rugosity at base of posterior USNM. horns. Anterior angles and fovea either side of longitudinal ridge rugose. Fovea deep. Horns: COLOMBIA (9).-CUNDINAMARCA (3): Bogota (?), Anterior short, stout, attenuate, curving forward Fusa(gasuga ?); SANTANDER (4): La Cimitarra and upward, apex narrowly rounded, dorsal sur­ (about 80 km. NW. Velez); No data (2). March (4), face flattened. Posterior horns very short, later­ May (2), October (1), November (1). ally compressed, reduced to large, prominent, Remarks.-The completely punctate meso­ subtriangular bosses; in lateral view dorsal sur­ sternum, lack of rows of punctures on the elytral face slopes backward and downward from a disc, presence of 2-3 short rows of moderate to rounded apex, anterior surface vertical; in dorsal large ocellate punctures behind the humerus, view horns divergent. Elytra: Sutural stria and the slightly expanded and weakly excised strongly impressed, crenulate. Disc aciculate, apex of the anterior horn will easily separate this sparsely punctate; punctures small and minute species. It is very closely related to S. jugurtha mixed, shallow; 3 shallow, incomplete striae on but cannot be confused with it because S. lateral half of disc; 5 large, ocellate punctures in jugurtha lacks the expanded and excised apex of a slight depression just medial of humerus at ex­ the anterior horn. S. fascinus is not commonly treme base. Sides similar except wrinkled, striae represented in collections and seems to be rarely absent; 2 very feeble rows of ocellate punctures collected. behind humerus; punctures sparse, moderate in size, shallow, becoming smaller posteriorly, a Strategus hipposiderus, new species few weakly umbilicate. Apex sparsely punctate; (Figs. 7, 9, 53-54, 138-139) punctures small, shallow. Pygidium: In lateral view basal half convex, apical half nearly flat. Type Material.-Holotype male, labeled "Val­ Disc finely subgranulate, aciculate, very sparsely ley of Rio Deseads, Nicaragua, April, 1931, C. D. punctate; punctures small, shallow, becoming 128 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

slightly more numerous near apical margin Females (11 paratypes): Length 47.7-55.6 mm; either side of midline; 1 setigerous puncture in a width across humerus 22.4-26.4 mm. Head: slight depression near center of disc. Apical Front rugose to rugo-punctate, setigerous above margin either side of midline with a very narrow, eye in unworn specimens. Clypeus with apex feeble, rugose band. Genitalia: Figs. 138-139. subacutely rounded to narrowly subtruncate, occasionally weakly emarginate, strongly re­ Allotype.-Female: Length 50.4 mm; width flexed. Interocular width 2.0-2.2 transverse eye across humerus 24.1 mm. As holotype except in diameters. Pronotum: Disc as allotype to moder­ the following respects: Head: Clypeus with apex ately densely punctate; minute punctures fewer, subacutely rounded; surface rugose. Tubercles several large punctures at midline. Elytra: Disc transverse. Mandibles with basal lobe as in with minute punctures very sparse, 3 feebly im­ ho~otype but slightly larger; middle lobe short, pressed striae on lateral half, 1-3 very short rows tri1¥1gular, apex rounded; apical lobe similar to of moderately large, ocellate punctures at ex­ basal lobe. Interocular width 2.0 transverse eye treme base just medial of humerus. Sides with . diameters. Pronotum: Disc sparsely punctate; 1-3 short rows of punctures; punctures as in al­ punctures small and minute mixed, shallow. lotype. Pygidium: Disc rugo-punctate to rugose, Sides and anterior half rugose; a slightly depres­ rarely sparsely punctate, sparsely setigerous in sed, rounded patch of rugosity just postero­ unworn specimens. Apical margin moderately lateral of fovea. Fovea deep. Tubercle strong, setigerous. conical, transverse. Elytra: Disc with punctures as holotype to slightly larger and denser; 2 very Biology.-Unknown. Label data indicate spec­ feebly impressed, incomplete striae on lateral imens have been taken at lights in Costa Rica half; several moderately large, ocellate, con­ and in the trunks of the coconut palm (Cocos fluent punctures just medial of humerus at ex­ nucifera L.) in Brazil. treme base. Pygidium: In lateral view basal half convex, apical half concave. Disc finely sub­ Distribution.-Southern Mexico to Panama, granulate, not aciculate, densely punctate; Colombia, and Brazil. The single specimen from punctures moderate to moderately large, round Brazil and the two records from Colombia estab­ to oblong, shallow. Apical margins either side of lish a very disjunct distribution. I believe the val­ midline rugose, sparsely setigerous; lateral idity of the Brazilian record is suspect. emargination shallow. Locality Records (Figs. 7, 9).-18 specimens Variation.-Males (4 paratypes): Length 48.0- examined (6 males, 12 females). Specimens are 54.0 mm; width across humerus 22.2-26.2 mm. deposited in the following collections: AHCC, Head: Clypeus as holotype to weakly rugulose. AMNH, BCRC, BMNH, DEUN, FMNH, HAHC, Pronotum: As holotype to sides with lateral mar­ USNM. gin rugose in a narrow band in basal half; a slightly depressed, rounded patch of rugosity BRAZIL (1 ).-BAHIA (1): Bahia. postero-Iateral of fovea. Anterior half rugose. COLOMBIA (2).-CAQUETA (1): Caucaya SE. Tres Horns: Anterior as holotype to very short, coni­ Esquinas on Rio Putumayo; No data (1). May (1), cal, erect. Posterior horns as holotype to very December (1). low, rounded bosses. Elytra: Disc and sides sub­ granulate, discal striae reduced to 2, rows of COSTA RICA (11).-LlM6N (10): Guapiles, Ham­ punctures behind humerus more distinct to vir­ burg Farm (Reventazon), Las Mercedes, Pan­ tually effaced. Pygidium: Disc with a shallow, dora; No data (1). March (1), May (2), June (2), median, longitudinal depression from center to August (3), September (1), December (1). apex; punctures becoming moderately dense MEXICO (1).-OAXACA (1): Juquila Mixes. July medially; 3-4 large setigerous punctures on disc; (1 ). apical band of rugosity slightly wider. Genitalia: Moderate variation in degree of arcuateness in NICARAGUA (1 ).-RIO SAN JUAN (1): Valley of Rio parameres. Deseads (sic). April (1). A REVISION OF THE GENUS Strategus I 129

PANAMA (2).-CANAL ZONE (2): Madden Dam. small, prominently rounded; middle lobe moder­ July (1), November (1). ate in size, subtriangular, apex rounded; apical lobe very small, rounded. Interocular width 2.66 Remarks.-The males of S. hipposiderus and transverse eye diameters. Mesosternum: An­ the larger, darker forms of S. aloeus are virtually terior half setigerously punctate, a few very identical externally, and examination of the sparse setae in posterior half. Pronotum: Base genitalia is required to separate them; fortu­ with a moderately wide rugose band; band re­ nately, the genitalia of the two species are vastly duced nearly to basal bead at midline. Disc different and cannot be confused. Difficulties aciculate, sparsely punctate; punctures small may also arise when separating males of S. hip­ deep. Sides in basal half as disc, puncture~ posiderus from the minor forms of S. man­ gradually becoming larger anteriorly; lateral dibularis. In general, however, S. mandibularis margin with a narrow rugose band; a slightly de­ minors have only a narrow to moderate basal pressed, rounded patch of rugosity on side of band of rugosity on the pronotum whereas it is posterior horns. Anterior half of pronotum always wide in S. hipposiderus; the punctures on rugose. Horns: Anterior very short, stout, at­ the disc of the mentum are usually large in S. tenuate, slightly transverse, apex subtruncate. mandibularis and small to moderate in S. hip­ Posterior horns are strong, laterally compressed posiderus, and there are usually some large bosses; in lateral view dorsal surface slopes for­ punctures on the disc of the pygidium in S. man­ ward and slightly downward, anterior surface dibularis and none in S. hipposiderus. In addi­ ~u.bvertical; in dorsal view horns parallel, bases tion, the parameres of S. hipposiderus are nor­ JOined across disc in an arc. Elytra: Sutural stria mally shorter and form a broader arc than do impressed, slightly wavy. Disc aciculate, sparsely those of S. mandibularis, but I have seen a few punctate; punctures small and minute mixed specimens of each where this distinction does shallow; small punctures umbilicate on lateral 213' not hold. The key characters will serve to sepa­ on median 113; 3 very feebly impressed: rate the females of S. hipposiderus, S. man­ ~imple Incomplete striae on lateral half. Sides wrinkled dibularis, and S. aloeus. behind humerus, otherwise as disc except striae Etymology.-From the Greek hipposideros, a absent and all small punctures umbilicate. Apex horseshoe; here named for the horseshoe­ densely punctate; punctures small and moderate shaped form of the male genitalia as seen in mixed, shallow. Pygidium: In lateral view basal caudal view. half convex, apical half nearly flat. Disc finely subgranulate, densely punctate; punctures small to large, shallow, occasionally confluent Strategus howdeni, new species setigerous apically either side of middle, seta~ (Figs. 6, 55-56, 140-141) small and fine. Genitalia: Figs. 140-141. Allotype.-Female: Length 37.7 mm; width Type Material.-Holotype male, labeled "Mex: across humerus 18.0 mm. As holotype except in N.L.; Monterrey, Chipenque Mesa, 5400', 22.vi. the following respects: Head: Front with supra­ ~969.' S. & J. Peck, forest litter, Berl 64"; depos­ ocular setae more pronounced. Clypeus with Ited In HAHC at CNCI. Allotype female with same apex narrowly truncate. Tubercles connected by data as holotype; deposited in HAHC at CNCI. a moderately strong, transverse carina. Pro­ Also one male paratype deposited in BCRC. notum: Disc with punctures slightly smaller. Holotype.-Male. Length 37.3 mm; width Tubercle strong, conical, transverse, apex trun­ across humerus 19.0. Color piceous, shining. cate. Elytra: Sutural stria crenulate. Disc with 2 Head: Front rugose, setigerous above eye. nearly obsolete striae and only a few scattered, Clypeus with apex rounded, strongly reflexed; umbilicate punctures. Pygidium: In lateral view surf.ace rugose. Tubercles moderately strong, basal half con"ex, apical half concave. Disc conical, transverse, weakly connected by a low, setigerous in a wide, irregular, median band; transverse carina. Mandibles with basal lobe setae very small, fine. 130 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Variation.-Single male paratype as holotype might occur between S. aloeus and S. howdeni except in the following respects: Length 38.0 as the latter were taken only near the summit of mm; width across humerus 19.3 mm. Head: In­ the mesa, but as of now I have no further infor­ terocular width 2.75 transverse eye diameters. mation to substantiate this. Pronotum: The slightly depressed area on side of Etymology.-This species is named in honor posterior horns sparsely punctate instead of of a friend and colleague, Henry F. Howden, who rugose, punctures moderate to large, shallow. collected the paratype example and in whose Horns: Posterior horns in lateral view with dorsal collection I first found specimens. edge-subhorizontal, anterior edge sloping back­ ward and upward; bosses not as developed as in holotype. Elytra: Sutural stria crenulate. Disc Strategus inermis Arrow with 2 feebly impressed striae and only a few (Figs. 2, 57-58, 142-143) scatlered umbilicate punctures. Pygidium: Sur­ f~ce aciculate, only moderately punctate. Strategus inermis Arrow, 1947: 223. [Holotype male at BMNH. Type locality: Haiti.] Biology.-Unknown. Distribution.-Chipenque Mesa, Monterrey, Male (Holotype).-Length 33.0 mm; width Mex;lpo. across humerus 15.5 mm. Color piceous, shin­ ing. Head: Front densely punctate to rugo­ Locality Records (Fig. 6).-3 specimens punctate, punctures large, shallow, confluent, examined (2 males, 1 female). Specimens depos­ setigerous above eye. Clypeus with apex broadly ited in BCRC and in HAHC at CNC!. truncate, moderately reflexed; surface densely MEXICO (3).-NuEVO LEON (3): Chipenque Mesa punctate; punctures large, shallow, confluent. (Monterrey). June (2), July (1). Tubercles conical, very feeble, transverse, widely separated. Mandibles with basal lobe feeble, Remarks.-The partially punctate mesoster­ hardly noticeable; middle lobe large, broadly num, distinctly lobed mandibles, the lack of rows rounded at nearly a right angle; apical lobe of moderate to large punctures on the elytral small, feeble, appearing as part of middle lobe. disc or behind the humerus, the shining elytra, Interocular width 2.33 transverse eye diameters. the simple and short anterior horn, the presence Mesosternum: Anterior half setigerously of a wide basal band of rugosity on the pro­ punctate. Pronotum: Base with a moderately notum, the rounded clypeal apex, the absence of wide band of dense punctures; punctures oval to strongly developed posterior horns, and the oblong, deep, some confluent. Disc very finely small to moderate punctures on the elytra should subgranulate, aciculate, sparsely punctate; serve to separate this species from all others. S. punctures small and minute mixed, shallow. howdeni seems to exhibit most of the external Sides in basal half as disc, lateral margin with a features of the Mexican population of S. aloeus moderately wide punctate band; punctures which is sympatric while the males possess moderately dense, large, shallow, some genitalia which are very similar to S. cessus cresent shaped, some confluent; a slightly de­ which is found further to the west. Strategus pressed, irregular rugo-punctate to punctate howdeni seems to be very much isolated in an patch postero-Iateral of fovea; punctures oak-pirie island mesa in a sea of desert scrub. cresent shaped, large, shallow. Anterior half In late June, 1974, I undertook a collecting trip rugose. Fovea shallow. Horns: Anterior a low, to Chipenque Mesa specifically to look for addi­ transverse, conical tubercle, apex emarginate. tional examples of S. howdeni. All levels of the Posterior horns nearly obsolete. Elytra: Sutural mesa were collected using blacklight and stria impressed, crenulate. Disc finely subgranu­ sodium vapor street lights. S. aloeus were found late, aciculate, with 8 feeble rows of punctures; in abundance (primarily at the lower levels of the punctures ocellate; small to moderate, shallow, mesa), but additional S. howdeni were not found. irregularly spaced; rows 3, 6, and 7 incomplete Quite possibly some sort of vertical isolation with punctures sparse, very small. Sides with 5 A REVISION OF THE GENUS Strategus / 131 ill-defined, confused rows of punctures;. prominently rounded; middle lobe moderately punctures ocellate, small to large, shallow. Apex large, triangular, apex rounded; apical lobe very densely punctate to rugo-punctate. Pygidium: small, triangular. Interocular width 2.0-2.66 Convex in lateral view. Disc finely subgranulate, transverse eye diameters. Mesosternum: Com­ densely punctate in a broad, irregular, median pletely and setigerously punctate. Pronotum: band; punctures setigerous, shallow, becoming Base with a narrow to obsolete rugo-punctate to reduced apically. Genitalia: Figs. 142-143. rugose band; band reduced to basal bead me­ dially. Disc aciculate, sparsely punctate; Female.-Unknown. punctures very small, deep, rarely with a few Biology.-Unknown. large punctures on midline. Sides as disc to punctures Slightly larger, lateral margin fre­ Distribution.-Haiti. quently with a rugo-punctate to rugose band in Locality Records (Fig. 2).-One male speci­ basal half; minors rugose in anterior half and men examined (holotype). Deposited at BMNH. with a slightly depressed, rounded patch of rugosity postero-Iateral of fovea. Anterior half as HAITI (1 ).-No data (1). disc in majors, usually rugose in minors. Fovea Remarks.-The partially punctate mesoster­ deep, median longitudinal ridge from base of an­ num, feebly lobed mandibles (as opposed to terior horn very low and broadly rounded, not sqdare in dorsal view), presence of rows of mod­ prominent. Horns: Majors (Figs. 59-60) with an­ erate to large punctures on the elytral disc, terior long, slender, attenuate, curving forward broadly truncate clypeal apex, shining elytra, ab­ and upward, apex narrowly rounded; dorsal sur­ sence of posterior horns, and the presence of a face rounded to nearly flat, when flat weakly and setigerously punctate band on the midline of the longitudinally carinate on lateral margins and pygidium will separate S. inermis from all other occasionally on midline. Posterior horns long, species. The holotype is the only known speci­ slender (slightly more robust than anterior horn). men of this species. Attenuate, laterally compressed, apex narrowly rounded, extending forward and upward at Strategus jugurtha Burmeister about 20-650 from plane of disc, not posteriorly (Figs. 7, 9, 59-62, 144-145) recurved apically; in dorsal view horns subparal­ lel to curved toward one another, bases joined Strategus jugurtha Burmeister, 1847: 131. [Lec­ across disc in a weak arc in all but the largest totype male and lectoallotype female (desig­ specimens, and then arc very indistinct. Minors nated by Endrodi, 1973a) at Martin Luther Uni­ (Figs. 61-62) with anterior as in major except versitat, Halle, German Democratic Republic. very short, dorsal surface rounded. Posterior Also two male and two female paralectotypes horns reduced to low pyramidal bosses; in lat­ at Halle. Type locality: Colombia.] eral view dorsal surface horizontal to sloping forward and downward, anterior surface subver­ Male.-Length 24.5-39.8 mm; width across tical to sloping upward and slightly backward; in humerus 12.2-18.5 mm. Color castaneous to dorsal view bosses laterally compressed, bases piceous, shining. Head: Front behind tubercles joined across disc in an arc. Elytra: Sutural stria strongly rugose in majors, less so to feebly impressed, crenulate. Disc finely subgranulate, rugose in minors; middle of front nearly im­ aciculate, sparsely punctate, punctures very punctate; setigerous above eye. Clypeus with small to minute, shallow, some occasionally apex acutely pointed to acutely rounded, occa­ raised; 1-3 feebly impressed striae on lateral half Sionally produced into an acute or rectangular present or not. Sides as disc except striae ab­ tooth, moderately reflexed; surface aciculate, sent, punctures slightly larger to moderate on weakly rugo-punctate to sparsely punctate; lateral margin; 2 (rarely 3) short rows of large punctures small to moderate, shallow. Tubercles punctures behind· humerus; punctures usually conical, moderately strong, usually distinctly ocellate, simple to umbilicate, deep. Apex separated. Mandibles with basal lobe small, sparsely to moderately punctate, punctures 132 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM SANTANDER (3): Bucaramanga, Soata, Velez; small to moderate, setigerous. Pygidium: Con­ VALLE (3): Anchicaya Dam (70 mi. ~. Buenaven­ vex in lateral view, apical half often nearly flat. tura); No data (1)). February (2), Apnl (1), May (3), Disc finely subgranulate, aciculate, very sparsely punctate; punctures minute or minute and mod­ June (3), July (2), December (2). erate mixed, shallow. Apical margin either side COSTA RICA (12).-GUANACASTE (,1).: Finca of middle with punctures slightly larger, or Jiminez (near Taboga); liM6N (9): Guaplles, Las weakly rugulose in a narrow band, not setiger­ Mercedes, Reventazon, Siguirres; No data (2). ous. Genitalia: Figs. 144-145. April (4), May (5), July (1), September (1), Female.-Length 25.01-36.5 mm; width across November (1). humerus 12.3-17.3 mnJ. "As male except in the ECUADOR (6).-PICHINCHA (3): Santo Domingo; following respects: H,e'ad: Front entirely rugose. SANTIAGO-ZAMORA (1): no data; No data (2). Au- Clypeus with apex 'C\:cutely rounded, more gust (5), October (1). broadly so if') worn specimens; surface weakly to EL SALVADOR (9).-SAN SALVADOR (3): San Sal­ strongly Fugose. TU,bercles transverse, fre­ vador; UNION (6): La Union, Vol. Conchagua. May quently connected by a leeble, transverse carina. Mandibles simila.,rto those of male except middle (9). lobe slightly smaller, apical lobe frequently a lit­ MEXICO (6).-CHIAPAS (1): Tuxtla Gutierre~; tle larger. Pronotum: Base usually with a narrow JALISCO (1): Casimiro Castillo; .SINALO.A (3): 6. ~I: rugose band; band reduced to basal bead at N. Mazatlan, Venedio, 28 mi. E. Villa Umon, middle. Sides with a slightly depressed, rounded VERACRUZ (1): Puente Nacional. June (3), July (2), patch of rugosity postero-Iateral of fovea. An­ August (1). terior half rugose. Pygidium: In lateral view basal NICARAGUA (6).-CHONTALES (4): no data; No half convex, apical half weakly concave to nearly data (2). July (1), September (1). flat. Disc finely subgranulate, virtually im­ punctate to moderately punctate; punctures PANAMA (60).-CANAL ZONE (38): Corozal, Cris~ small to moderate, shallow, setigerous in un­ tobal, Las Cascadas, Madden ~am.; DARIEN (16). worn specimens. Lateral emargination very shal­ Sante Fe' PANAMA (2): Panama City; VERAGUAS low to obsolete, occasionally weakly rugose. (1): Santi'ago; No data (3). May (23), June (31), October (1 ). Biology.-Basically unknown. Lamb (1974) re­ ported that S. jugurtha attacks pineapple stems PERU (1 ).-LORETO (1): Varina Cocha (Ucayali in Central and South America. Label data indi­ River). cate that this species has been widely taken at Remarks.-Strategus jugurtha can usually be lights, from agave in Colombia, pineapple in quickly identified by the sh~rt s~tae on the Costa Rica, and attacking the roots of African oil apices of the elytra in combination With the large palm (Elaeis guineensis Jacq.) in Nicaragua. punctures behind the humerus. If the seta~ are Distribution.-Mexico through Central worn off, this species may b~ con~used With S. America to Colombia west of the Cordillera surinamensis surinamensis, m which case the Orienta and Peru. The single Peruvian record is key characters and the place of origin shou~d be questionable. sufficient to separate the two. The Cordl.llera Orienta in northern Colombia and the Cordlll~ra Locality Records (Figs. 7,9).-126 specimens do Merida in western Venezuela seem t~ prOVide examined (60 males, 66 females). Specimens a distinct barrier between the two species. were seen from the following collections: AHCC, AMNH, BMNH, CASC, CNCI, FMNH, HAHC, LACM, MCZC,PMNH, UNC,USNM. Strategus longichomperus, new species (Figs. 7,63-66, 146-147) COLUMBIA (26).-BoYACA (1): Miraflores; CUN­ Type Material.-Holotype male, labeled "San DINAMARCA (2): Bogota; HUILA (3): Gigante, Andres, 6/30/61" (= San Andres Tuxtla, Vera- Neiva; MAGDALENA (2): Aracataca; NORTE DE A REVISION OF THE GENUS Strategus / 133

cruz, Mexico); deposited at FMNH. Allotype cally either side of midline, a few scattered, female, labeled "Guatamala: Peten, Tikal, 18Apr. large, shallow punctures medially. Apex with 1956, at camp., E. M. Shock"; deposited at several moderate-sized, setigerous punctures; FMNH. Also six male and seven female paratypes setae long. Genitalia: Figs. 146-147. deposited in the following collections: AMNH, Allotype.-Female. Length 41.7 mm; width BCRC, BMNH,. CNCI, MCZC, SEBD ENDRODI, across humerus 19.5 mm. As holotype except in USNM. the following respects: Color more reddish. Head: Front completely rugose. Clypeal apex Holotype.-Male. Length 45.6 mm; width with a small, triangular notch; surface rugose. across 22.4 mm. Color piceous, shin­ humet~~ Mandibles similar to those of holotype but smal­ ing. Head: I7liant behind each tubercle densely ler, especially middle lobe; middle lobe with punctate; p.un~tures large, shallow, some con­ apex acutely rounded. Pronotum: Disc very fluent; middle"of front virtually impunctate, sur­ sparsely punctate; punctures small to minute. fas;:e Clypeus with apex strongly re­ roughen~d. Sides rugose; a slightly depressed, rounded flexed, deeply ~rd broadly excised by a triangu­ patch of rugosity postero-Iateral of fovea. An­ lar notch, I,obes either side of emargination terior third rugose. Fovea deep. Tubercle very triangular; surface aciculate, sparsely punctate; low, conical, transverse. Elytra: Disc with punctures moderately deep, very shallow. punctures very small, extremely shallow to obso­ Tubercles moderately strong, conical, trans­ lete, some almost in rows. Pygidium: In lateral verse, weakly separated. Mandibles with basal view basal half convex, apical half concave. Disc lobe small, rounded; middle lobe large, elongate, finely subgranulate, densely and completely triangular, apex rounded; apical lobe small, setigerously punctate; punctures moderate to triangular. Interocular width 2.25 transverse eye large, shallow, setae long; lateral emargination diameters. Mesosternum: Anterior half setiger­ very shallow. ously punctate. Pronotum: Base with a narrow, rugose band; band reduced to basal bead at Varlation.-Males (6 paratypes): Length 35.4- middle. Disc aciculate, sparsely punctate; 48.4 mm; width across humerus 16.3-23.3 mm. punctures small, deep. Sides as disc except lat­ Head: Front as holotype to rugose in minors. erm margin in basal half with a few large, shallow Clypeus as holotype except minors with apex punctures. Anterior half as disc. Fovea deep, weakly and broadly emarginate; surface rugose. median longitudinal ridge from base of anterior Interocular width 3.0 transverse eye diameters in horn very low and broadly rounded, not promi­ minors. Pronotum: As holotype except minors nent. Horns: Anterior long, subslender, at­ with a narrow, punctate to rugo-punctate band tenuate, curving forward and upward, apex nar­ on lateral margin in basal half. Anterior half rowly rounded, dorsal surface rounded. Poste­ rugose. Horns: Majors (Figs. 63-64) with anterior rior horns moderate in length, attenuate, robust, as holotype to apex weakly and minutely emar­ laterally compressed, projecting forward sub­ ginate in 1 paratype. Minors (Figs. 65-66) with horizontally, apex rounded; in lateral view horns anterior a strong, transverse, conical tubercle. parallel, bases joined across disc in an arc. Posterior horns virtually effaced. Elytra: Disc Elytra: Sutural stria strongly impressed, crenu­ frequently with 2 feebly impressed, incomplete late. Disc aciculate, sparsely punctate; striae. Sides with a short, median, third row of punctures small, deep. Sides as disc except ocellate-umbilicate punctures; minors with lat­ punctures slightly larger, wrinkling behind eral rows of ocellate-umbilicate punctures nearly humerus, and with 2 short rows of ocellate­ obsolete. Pygidium: Disc on midline with an ir­ umbilicate punctures behind humerus; regular, longitudinal, narrow to wide band of punctures large, shallow, median row forked. setigerous punctures, setae long; remainder of Apex sparsely punctate; punctures small, deep. disc without setigerous punctures. Apical mar­ Pygidium: Convex in lateral view. Disc finely gins occasionally rugo-punctate to rugose in a granulate, aciculate, moderately punctate; wide band, setigerous. punctures small, shallow, becoming denser api- Females (7 paratypes): Length 37.4-43.1 mm; 134 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM width across humerus 17.0-20.5 mm. Color as erence to the very elongate middle lobe of the holotype to more reddish as allotype. Head: mandibles. Front densely punctate, punctures large, shal­ low, confluent. Mandibles as allotype to apex of middle lobe not as acute, more rounded. In­ Strategus mandibularis Sternberg terocular width 2.0-2.5 transverse eye diameters. (Figs. 9, 67-70, 148-149) Pronotum: Base with rugose band very narrow to obsolete. Sides with basal half as disc except Strategus mandibularis Sternberg, 1910: 99. punctures slightly larger and denser, a narrow [Holotype male at ZMHU. Type locality: Montes rugose band on lateral margin. Tubercle very low Claros, Minas Gerias, Brazil.] and rounded to strong and conical. Elytra: Disc with punctures minute to small, very shallow to Male.-Length 51.4-57.5 mm; width across moderat~ly deep, occasionally with 2 feebly im­ humerus 24.0-27.0 mm. Color piceous, shining. pressed, incomplete f?triae. Sides as allotype to Head: Front grossly rugo-punctate to rugose, rows of ocellate-umbilicate punctures obsolete weakly setigerous above eye in unworn speci­ in largest specimens. Pygidium: Disc with mens. Clypeus with apex broadly truncate, punctures reduced to moderate in density in strongly reflexed; surface rugulose to coarsely largest specimens. punctate; punctures large, confluent, and shal­ low, or sparsely punctate; punctures small, deep. Biology.-Un known. Tubercles conical, moderately strong, trans­ verse, distinctly (but not widely) separated. Man­ Distribution.-Central America. dibles with basal lobe small, rounded; middle Locality Records (Fig. 7).-15 specimens lobe large, triangular, apex rounded; apical lobe examined (7 males, 8 females). Specimens de­ small, triangular. Interocular width 2.5 transverse posited in the following collections: AMNH, eye diameters. Mesosternum: Anterior half BCRC, BMNH, CNCI, FMNH, MCZC, SEBO EN­ setigerously punctate. Pronotum: Base with a DROOl, USNM. wide band of rugosity; band reduced to basal bead at middle. Disc aciculate, feebly roughened GUATAMALA (5).-EL PETEN (3): Tikal; No data to finely granulate, sparsely punctate; punctures (2). April (3), May (1). small, deep. Sides similar except punctures a lit­ HONDURAS (2).-ATLANTIDA (2): La Ceiba. April tle larger and denser laterally; lateral margin with (1), July (1). a narrow punctate to rugo-punctate band in basal %; base of posterior horns usually with an MEXICO (8).-CHIAPAS (2): Palenque; irregular, slightly depressed, punctate to rugo­ TAMAULIPAS (3): Ciudad Victoria, EI SaIto Falls punctate area. Anterior third as disc in majors, (26 mi. W. Antiguo Morelos); VERACRUZ (3): Dos rugose in minors. Fovea shallow, median lon­ Amates (Catemaco), Lake Catemaco, San Andres gitudinal ridge from base of anterior horn strong, Tuxtla. May (1), June (3), July (1), August (3). rounded to keel-like. Horns: Majors (Figs. 67-68) Remarks.-The key characters will serve to with anterior long, slender, attenuate, curving separate S. longichomperus from all other forward and upward, apex narrowly rounded; Strategus, although the deeply excised clypeal dorsal surface rounded to subcarinate. Posterior apex, the very elongate mandibles, and the sub­ horns long, slender, attenuate, laterally com­ vertical posterior horns in the males will be im­ pressed, extending forward and upward at about mediately noticed. S. longichomperus is the only 65-75° from plane of disc, apex narrowly species in the genus where the female possesses rounded; in dorsal view horns subparallel to a triangularly notched clypeal apex. strongly curving toward one another, bases not forming a distinct arc across disc. Minors (Figs. Etymology.-From the Latin longus, meaning 69-70) with anterior similar to majors but very long, and from the American slang "chompers" short. Posterior horns reduced to prominent, meaning teeth; here descriptively named in ref- triangular bosses. Elytra: Sutural stria strongly A REVISION OF THE GENUS Strategus / 135 impressed, crenulate. Disc aciculate, sparsely seen from the following collections: UC, USP, punctate, punctures small and minute mixed, ZMHU. very shallow to deep, occasionally some slightly ARGENTINA (2).-MISIONES (2): Loreto. De­ raised; 2-3 very feebl.y impressed striae on lateral cember (2). half present or not. Sides similar except 2-3 very weak rows of moderate sized, shallow punctures BRAZIL (5).-GOIAS (2): 20 km. N. Sao Joao da behind humerus, rpws occasionally effaced; Alianca; MINAS GERIAS (1): Montes Claros; SANTA striae lacking. Ap~~ moderately punctate; CATARINA (1): Rio Vermhelho; SAo PAULO (1): punctures small to;mOderate, shallow. Pygidium: Ararquara. March (1), April (1), May (2). Convex in lateral vieW. Disc finely subgranulate, PARAGUAY (2).-BOQUER6N (1): Gran Choco; aciculate, sparsely pimctate; punctures minute GUAIRA (1): Puerto Suayra (Villarrica). June (1). to small" shallow; oqcasionally a strong, median band of large pun'~~res present; punctures Rernarks.-The male of S. mandibularis and S. dense, oval to oblong, shallow, setigerous. Api­ validus are similar, but S. mandibularis can be cal margins either side of middle rugo-punctate quickly separated by its large, middle mandibu­ in a narrow band to moderately densely lar lobe, the wide band of rugosity at the base of punctate; punctures small to moderately large, the pronotum, and the usually more punctate shallow, simple to oblong, sparsely setigerous in pygidium. Male minors of S. mandibularis might unworn specimens; setae long. Genitalia: Figs. be confused with S. hipposiderus, but the '148-149. genitalia of each are diagnostic. See also the re­ marks for S. hipposiderus. The key characters Fernale.-Length 51.2-52.2 mm; width across will adequately separate the females. The humerus 24.3-24.7 mm. As male except in the females up until this time were apparently un­ following respects: Head: Clypeus with apex nar­ known and are here recogn ized and described rowly truncate; surface rugose. Mandibles simi­ for the first time. lar to those of male but smaller, especially mid­ dle lobe. Interocular width 2.75 transverse eye diameters. Pronotum: Base with rugose band Strategus mormon Burmeister narrow to wide. Sides rugo-punctate to rugose in (Figs. 6, 71-72, 150--151) basal third, anterior 213 rugose; a slightly depres­ sed, rounded patch of rugosity postero-Iateral of Strategus mormon Burmeister, 1847: 130. fovea. Anterior half rugose. Fovea moderately [Holotype male at MNHN (Oberthur Collec­ deep. Tubercle conical, strong, slightly trans­ tion). Type locality: Texas.) verse. Elytra: Disc at base just medial of humerus with 1-4 very short, confused rows of punctures; Male.-Length 25.0-29.2 mm; width across punctures ocellate-umbilicate, small to large, humerus 13.6-15.8 mm. Color dark castaneous, dense, shallow. Sides behind humerus usually occasionally piceous, shining. Head: Front with 4 short, irregular rows of punctures; rugo-punctate to coarsely punctate; punctures punctures ocellate-umbilicate, small to large, small to moderately large, dense, shallow, con­ shallow; rows occasionally reduced or obsolete. fluent; setigerous above eye. Clypeus with apex Pygidium: In lateral view basal half convex, api­ very acutely pOinted; surface aciculate, weakly cal half concave. Surface entirely rugose, den­ rugo-punctate to densely punctate; punctures sely setigerous. small to moderately large, shallow, becoming Blology.-Unknown. smaller and denser laterally. Tubercles conical, transverse, very weak to virtually effaced. Man­ Distribution.-Argentina, Brazil, and dibles with basal lobe small, rounded; middle Paraguay. lobe very long, slender, apex acutely pointed; apical lobe small, triangular. Interocular width Locality Records (Fig. 9).-9 specimens 3.0--3.5 transverse eye diameters. Mesosternum: examined (4 males, 5 females). Specimens were Completely and setigerously punctate. Pro- 136 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

notum: Base with a very narrow, sparsely or rugose in apical third. Fovea shallow. Tuber­ punctate band; punctures small to moderately cle conical, weak to strong, transverse, apex large, shallow, oval to oblong; band reduced usually weakly emarginate. Pygidium: In lateral nearly to basal bead at midline. Disc aciculate, view basal half convex, apical half concave. Disc sparsely punctate; punctures small and minute with punctures occasionally large. Apical mar­ mixed, shallow. Sides similar to disc except lat­ gins moderately to densely punctate; punctures eral margin with a narrow band of sparse simple to oblong, shallow. punctures; punctures small to moderately large, Biology.-Knaus (1916) observed S. mormon shallow, occasionally setigerous at middle in burrows under and adjacent to horse droppings unworn specimens, becoming rugose in anterior in Kansas and indicated that burrow shafts were angles; a slightly depressed, rounded, punctate vertical or inclined up to 15° and that they varied area postero-Iateral of fovea; punctures small to from 4-12 inches in depth. A mass of dung ,:moderate, dense to nearly obsolete, shallow. An­ 1.25-1.50 inches in diameter and several inches terior margins rugose, rugosity frequently ex­ long was found in a burrow under 3-4-week-old tending postero-medially into fovea. Fovea mod­ horse manure; seven white eggs measuring 2-3 erately deep, only occasionally weakly divided by mm in diameter were found in the dung. Other a feeble, longitudinal, median carina extending similar burrows were found by Knaus who "posteriorly from base of anterior horn. Horns: suggested that this species prefers horse drop­ anterior conical to very short, slender, attenuate, pings 1-3 weeks old; S. mormon burrows were transverse, curving forward and upward, apex never associated with cow manure although narrowly rounded to minutely emarginate. Pos­ Knaus later mentions that he did find a dead terior horns are weak to very prominent triangu­ female with an associated burrow under cow lar bosses; in lateral view dorsal edge slopes manure. In a later paper Knaus (1924) concluded, forward and downward to apex in undeveloped after examining many burrows, that the male ap­ specimens, and backward and downward from pears 5-15 days before the female and prepares apex in developed specimens, anterior edge a burrow under horse manure, preferably select­ slopes backward and upward in undeveloped ing droppings 6-12 months old. The bottom of specimens and is subvertical in developed indi­ the burrow is enlarged to contain the larval pro­ viduals; in dorsal view bosses laterally compres­ visions which consist of horse or cow manure sed, parallel, bases joined across disc in an arc. obtained by the attracted female. Several eggs Elytra: Sutural stria weakly to moderately are deposited in the food mass after which the impressed, subcrenulate. Disc aciculate, very adults leave the burrow. If true, the apparent di­ sparsely punctate; punctures minute and small vision of labor and burrow prOVisioning de­ mixed, shallow; 2-3 very feebly impressed, in­ scribed (or generalized upon) by Knaus would be complete striae on lateral half, striae subgranu­ remarkable for dynastine scarabs which are not late. Sides as disc except striae absent. Apex de­ generally known for such behavior. More de­ nsely punctate; punctures small to moderate, tailed observations are certainly needed to verify shallow, with very short setae in unworn speci­ Knaus' conclusions. mens. Pygidium: Convex in lateral view. Disc Strategus mormon seem to be primarily finely subgranulate, very sparsely punctate; arenicolous. This species is apparently relatively punctures moderate in size, very shallow. Apical scarce in numbers, and field collecting and margins either side of middle with punctures a museum collections tend to support this view. In little larger, denser, and deeper. Genitalia: Figs. two days of very intense collecting at Medora, 150--151. Kansas, in late May, 1972, I was able to obtain only one specimen at blacklight even though Female.-Length 26.0--28.8 mm; width across horse and cattle droppings were examined in humerus 13.5-15.0 mm. As male except in the large numbers. following respects: Pronotum: Apical half mod­ erately to densely punctate; punctures small to Distribution.-Southcentral United States. large, shallow, usually grading to rugo-punctate Locality Records (Fig. 6).-186 specimens A REVISION OF THE GENUS Strategus / 137

examined (91 males, 95 females). Specimens coarsely rugo-punctate to rugose, occasionally were seen from the following collections: AMNH, very reduced in especially large individuals. BCRC, CASC, CNCI, DE UN, FMNH, HAHC, KSUC, Clypeus with apex strongly reflexed, very broadly LACM, MCZC, OSUC, OSUO, PMNH, SEMC, truncate, frequently with a very shallow, triangu­ UMMC, USNM. lar emargination; surface in majors punctate; punctures moderately dense, small, shallow; sur­ UNITED STATES (186).-ARKANSAS (2): Benton face in minors coarsely rugo-punctate to rugose. (1): Bentonville; Pope (1): Russelville. KANSAS Tubercles conical, low, transverse, usually dis­ (142): McPherson (2): 18 mi. S. McPherson; tinctly separated. Mandible with basal lobe Remo (129): Medora, Sylvia; Rice (1): 8 mi. S. lit­ small, prominently rounded; middle lobe large, tle River. OKLAHOMA (39): Delaware (1): Jay; triangular, apex rounded; apical lobe small, sub­ Ki(1gfisher (1): Kingfisher; Mayes (2); Chatou; triangular. Interocular width 2.0-2.5 transverse Murray (1): Sulphur; Muskogee (2): Muskogee; eye diameters. Mesosternum: Anterior half ()klahoma (1): Norman; Pawnee (4): no data; setigerously punctate. Pronotum: Base with a 'Payne (23): Stillwater; Pottawatome (2): Shaw­ narrow to moderately wide rugose band; band nee; Tulsa (1): Tulsa; No data (1). TEXAS (3): reduced to basal bead at midline. Disc aciculate, .Bexas (1): San Antonio; No data (2). March (1), sparsely punctate; punctures small, deep. Sides '~pril (1), May (29), June (134), July (11). I, similar except minors usually with a feeble, Remarks.-This species is easily recognized sparsely punctate band on lateral margin; by its very long, slender, acute middle lobe of the punctures large, round to oblong. Anterior por­ mandibles in conjunction with a completely, tion of pronotum as disc in majors; minors with setigerously punctate mesosternum. The males anterior Y4-Y3 rugose or rugose in a band from do not display what could be accurately termed anterior angles postero-medially into fovea. major and minor development of pronotal arma­ Fovea deep; longitudinal median carina extend­ ture although minor differences in horn length ing posteriorly from base of anterior horn nearly do exist. obsolete. Horns: Majors (Figs. 73--74) with an­ terior long, subslender, attenuate, curving for­ ward and upward, apex narrowly rounded; dor­ Strategus ob/ongus (Beauvois) sal surface weakly convex to flattened. Posterior (Figs. 2,73--76,152-153) horns long, stout, attenuate, laterally compres­ sed, extending forward and upward at about Scarabaeus oblongus Beauvois, 1807: 74. [Types 25-45° from plane of disc, apex narrowly unknown to me. Type locality: Dominican Re­ rounded; in dorsal view horns parallel or curving public.] toward one another, bases weakly joined across Scarabaeus quadrifoveatus Beauvois, 1807: 74. disc in an indistinct arc. Minors (Figs. 75-76) [Types unknown to me. Type locality: Domini­ with anterior as majors except short, stout. Pos­ can Republic.] terior horns nearly obsolete to prominent, acute, Scarabaeus beauvoisi, New name. [Replaces laterally compressed, pyramidal bosses; the lat­ Scarabaeus dubius Beauvois, 1819: 209, which ter in lateral view with dorsal surface sloping is a primary junior homonym with Scarabaeus forward and downward to apex or backward and dubius Beauvois, 1811: 104 (a Dyscinetus sp.) downward from apex, anterior surface vertical to which, in turn, is a primary junior homonym subvertical; in dorsal view horns subparallel, with Scarabaeus dubius Olivier, 1789: 32 (also bases joined across disc in an arc. Elytra: Sutural a Dyscinetus sp.). Types unknown to me. Type stria strongly impressed, distinctly crenulate to locality: Dominican Republic.] New wavy. Disc aciculate, finely subgranulate, synonymy. sparsely punctate; punctures small and minute mixed, deep, rarely slightly raised; usually 2 fee­ Mate.-Length 35.0-58.8 mm; width across bly impressed, incomplete striae on lateral half. humerus 18.0-27.8 mm. Color piceous to black Sides as disc except punctures slightly larger to (rarely castaneous), shining. Head: Front moderate in size, frequently a short row of 1-10 138 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

moderate to large, very shallow punctures be­ concealed in small cases of wood and humus. hind humerus, and striae absent. Apex moder­ Wolcott (1948) noted that the eggs are also laid ately punctate; punctures small, deep. Pygidium: in the rotting trunks of dead palms which quickly In lateral view basal 3/4 convex, apical 1/4 concave. collapse after the larvae feed on the interior. The Disc finely subgranulate, sparsely to moderately larvae are about 10 mm in length when they densely punctate; punctures small to moderate, hatch and have been seen to .consume their egg shallow when sparse, very shallow to nearly ef­ shells and to eat cast skins. Tables 1 and 2 illus­ faced when dense. Apical margins rugose either trate larval development. side of middle; lateral emargination shallow. Genitalia: Figs. 152-153. TABLE 1 F~male.-Length 39.4-48.2 mm; width across INSTAR DURATION IN DAYS FOR S. obJongus hurii.~tus 18.3-22.3 mm. As male except in the following respects: Head: Front rugose, setiger­ iNSTAR No. OesERvED RANGE AVERAGE ous above eye in unworn specimens. Clypeus 48 with"apex broadly truncate to narrowly rounded. 46-144 73 2 9 90-240 145 Ma(ldibles similar to those of male but smaller. 3 6 180-240 210 Pro~qtum: Base with a very narrow rugose band; band reduced at middle almost to basal bead . .... TABLE 2 Sides with a narrow rugo-punctate to rugose INSTAR MEASUREMENTS IN MM FOR S. obJongus band on lateral margin in posterior half; a simi­

lar, but very wide, band of rugosity in anterior HEAD CAPSULE WIDTH iNSTAR No. OesERvED MAXIMUM BODY LENGTH half; frequently a small, slightly depressed, RANGE AVERAGE punctate to rugose patch lateral of base of fovea; punctures moderate to large, sparse to dense, 1 296 2.8-4.0 3.6 28.0 round to oblong to crescent shaped. Anterior 2 176 5.&-7.7 6.7 55.0 3 249 9.6-14.1 12.3 100.0 third rugose. Fovea moderate in depth. Tubercle conical, transverse, weak to moderately strong. Pygidium: In lateral view basal half convex, api­ cal half concave. Apical margins with a wide, Larvae became cannibalistic when crowded rugose band either side of middle. and were also seen to chew through the bottom Biology.-Plank (1945), in a verification and of tin-can rearing containers, thus giving some supplement to Smythe (1916, 1920), has given a idea as to their mandibular strength. The larvae detailed account of the life history of S. oblongus feed on the roots of deciduous trees and in in Puerto Rico as follows: The eggs are large moist, decaying tree stumps (Wolcott, 1949). (2.9-3.1 x 3.7-4.0 mm), globular to oblong, with Larvae have been taken from the following food a smooth chorion, and are an opaque, ivory hosts: Cocos nucifera L., Citrus sinensis (L.), white when newly deposited. During incubation Inga inga (L.), Inga laurina (Sw.), Spathodea they become somewhat translucent and after 10 campanulata Beauv., Spondias sp., and other days assume a pearly white color which later palms. They have also been observed damaging changes to dull ivory. Eggs may expand while the roots of young cacao'trees (Theobroma incubating; the largest egg seen was 4.5 x 5.4 cacao L.) in the Dominican Republic (Wolcott, mm two days before hatching. A total of 51 eggs 1926). Under certain conditions larvae fed on the were observed to have an incubation period of rind of decaying coconut husks but not on the 13-34 days (average 16.8 days). Eggs with the fibers. Larvae were unable to survive on sugar shortest incubation periods were deposited in cane and old manure (Smythe, 1920). June and July; those with the longest were laid in The pre-pupal state in 18 specimens lasted February when temperatures were cooler. from 12-20 days (average 18 days). The pupal The eggs may be scattered in the softer, de­ period in 60 specimens ranged from 20-47 days cayed parts of loose wood but are more usually (average 33 days). Newly emerged adults were A REVISION OF THE GENUS Strategus / 139 observed to remain in the pupal chamber 2-10 several individuals may congregate on one stool days (average four days) after eclosion. while those nearby are unharmed. Lodging may The length of adult life ranged from 53 to 193 occur when a beetle bores into the stalk at days (average 90 days) in 63 beetles. Adult male ground level and tunnels upward 8-14 inches. longevity was 61-128 days (average 96 days) in The attempted use of dead palms for fence posts 29 specimens. Adult female longevity was 53-131 or anything else is impossible and actually days (average 85 days) in 34 beetles. The total life hazardous to the palm groves because they may span in S. ob/ongus may be as long as 16-18 contain developing beetles within the palm logs. months . S. ob/ongus may become injuriously abundant a . 'Plank noted that eggs were deposited nine few years after destructive hurricanes unless the days after the one observed mating, and oviposi­ dead palms are cleared away (Wolcott, 1948). tion continued for nine days; 10 eggs were de­ The regions of greatest wind damage (not posited, and four well-developed eggs remained necessarily those of the greatest coconut pro­ hi the female when she died. All stages, except duction) have usually been the areas suffering the egg, have been found in all months of the the most insect injury. year. The severe cutting of the native forests has Feeding habits and gut analyses demonstrated driven Strategus to live on sugar cane and palm that the adults feed primarily on the juices of the trees and to become economically harmful. Wol­ pf~nts they attack. Martorell (1945a) and Wolcott cott (1936) lists several economic accounts and (1923, 1948) observed that adults occasionally illustrates the larvae. burrowed into the stalks of sugar cane and in­ jured young coconut palms by burrowing into Distribution.-Grand Cayman Island, His­ the trunk and roots at ground level. paniola, Puerto Rico; not established on St. Natural enemies are apparently few in number. Croix (Miskimen and Bond, 1970). The short Nematodes attacked the eggs in the laboratory series of specimens from Grand Cayman is par­ but were not seen in the field. The green mus­ ticulary interesting because they are reasonably cardine fungus, Metarrhizium anisop/iae disjunct in their distribution. The prevailing (Metschn.), attacked some larvae. Martorell winds, currents, and storm paths, however, (1945b) observed that both the adults and imma­ might easly account for establishment of S. ob­ ture stages are susceptible to attack by the fun­ longus on Grand Cayman from a Hispaniola gus with the pupa being the most susceptable source. and the egg the least susceptible to attack. Pre­ dators include mites of the families Parasitidae Locality Records (Fig. 2).-175 specimens and Tyroglyphidae as well as herons, owls, rats, examined (75 males, 100 females). Specimens poultry, hogs, and the mongoose. Martorell were seen from the following collections: AMNH, (1945b) noted that the mongoose, Herpestes BCRC, BMNH, CNCI, HAHC, MCZC, TAMU, birmanicus (Thomas), which feeds on the adults, USNM. might be the most important enemy of S. ob/on­ GRAND CAYMAN ISLAND (5).-GRAND CAYMAN gus, while the Puerto Rican grackel, Quisca/us (5): West end of Georgetown. May (1), July (4). niger brachypterus (Cassin), is very efficient in consuming larvae when they are turned by plows DOMINICAN REPUBLIC (41).-DISTRITO NA­ in fields under cultivation. ClONAL (1): Santo Domingo; LA VEGA (9): Con­ Known as the coconut rhinoceros beetle, S. stanza, Convento (12 km. S. Constanza), ob/ongus was a serious pest of coconuts in Jarabacoa, Rio Camu (19 km. NE. Jarabacoa); Puerto Rico (Martorell, 1945a; Plank, 1945) but MONTE CRISTI (1): Monte Cristi; PERAVIA (3): was insignificant as a pest of sugar cane Bani; PUERTO PLATA (2): Puerto Plata, Sosua; (Smythe, 1920). The principal damage results SAMANA (9): Sanchez; SAN PEDRO DE MACORIS from adults feeding on germinal tissue and the (5): San Pedro de Macoris; SANTIAGO (2): San­ growing point of palms. The adults have been tiago; No data (9). May (4), June (15), July (6), taken a number of times injuring sugar cane; August (7), October (2). 140 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

HAITI (40).-OUEST (38): Port-au-Prince; No data above eye in unworn specimens. Clypeus with (2). February (2), March (1), May (2), August (1), apex rounded, moderately reflexed; surface October (31). rugo-punctate to weakly or moderately punctate; punctures confluent, shallow. Tubercles conical, PUERTO RICO (89).-AGUADILLA (6): Aquadilla, very low to nearly effaced, transverse, usually Anasco, Lares, Las Marias; ARECIBO (4): Arecibo, widely separated although occasionally con­ 5 mi. NE. Jayuya, Manati, Utuado; GUAYAMA (3): nected by a feeble, transverse carina. Mandibles Aguirre, Aibonito, Caguas; HUMACAO (4): EI Yun­ with basal lobe small, rounded; middle lobe que Sta. (Luquillo Forest), Loiza Viejo, Luquillo, moderate in size, subobtusely to obtusely Rio Grande; MAYAGUEZ (52): Cabo Rojo, rounded; apical lobe very small, broadly con­ Maricao, Mayaguez, San German, Yauco; PONCE fluent with middle lobe. Interocular width about (2): Adju.ntas, Ponce; SAN JUAN (5): Bayamon, 3.0 transverse eye diameters. Mesosternum: An­ Rio Piedras, San Juan, Toa-Baja; No Coro~ar,·· terior half setigerously punctate. Pronotum: data (JS). January (6), February (6), March (4), Base with a moderately wide, rugo-punctate to April ~(~), May (6), June (5), July (7), August (5), rugose band; band reduced nearly to basal bead Septe'r;'nber (13), October (10), November (1), De­ medially. Disc aciculate, sparsely punctate; cember (12). punctures small, moderately deep. Sides similar REMARKS.-The broadly truncate clypeus, to disc except punctures slightly larger; lateral narroWiband of rugosity on the base of the pro­ margin with a moderately wide, rugo-punctate to : notum, simple horns, and the lack of large, rugose band joined to basal band; frequently a post-humeral or discal punctures will usually rounded, slightly depressed area of moderate to separate S. ob/ongus from all other species. large punctures just postero-Iateral of fovea. An­ Arrow (1937a), in synonymizing S. quad­ terior %-Y3 rugose. Fovea shallow. Horns: An­ rifoveatus with S. ob/ongus, was apparently not terior a moderate to large, slightly transverse satisfied that S. quadrifoveatus was a Strategus. tubercle; apex narrowly rounded or narrowly Indeed, Beauvois' illustration depicts a specimen truncate. Posterior horns completely lacking. lacking a fovea on an otherwise convex pro­ E/ytra: Sutural stria impressed, usually not cre­ notum; in addition, the elytra possesses 2-3 nulate. Disc aciculate, medial Y3-Y2 sparsely to striae. Without the type it is difficult to state posi­ moderately punctate; punctures ocellate­ tively that S. quadrifoveatus was a Strategus, but umbilicate, moderate to large, shallow; lateral it will be so considered here. It is definitely Y~/a usually with 6 rows of moderate to large, strategid in its affinities and may simply be a ocellate-umbilicate, shallow punctures, rows 1-2 female with a nearly obsolete pronotal fovea. and 4-5 closely adjacent, rows 3 and 6 sparsely Although S. quadrifoveatus was described punctate; rows occasionally reduced. Sides usu­ prior to S. ob/ongus, it is here formally desig­ ally with 6 confused rows of punctures similar to nated that Strategus ob/ongus is the valid name those of disc. Apex moderately densely for this taxon in order to maintain nomenclatural punctate; punctures small and moderate mixed, stability. shallow. Pygidium: In lateral view basal half strongly convex, apical half weakly concave to Strategus sarpedon (Burmeister) nearly flat. Disc very finely granulate, sparsely to (Figs. 2,77-78,154-155) moderately punctate; punctures small, very shal­ low; disc occasionally becoming weakly rugo­ Poda/gus sarpedon Burmeister, 1847: 122. punctate. Apical margins frequently rugo­ [Types unknown to me. Type locality: Cuba.] punctate to rugose. Genitalia: Figs. 154-155.

Male.-Length 19.9-28.2 mm; width across Female.-Length 21.2-25.0 mm; width across humerus 10.4-13.9 mm. Color castaneous to humerus 10.9-12.5 mm. As male except in the piceous, shining. Head: Front grossly punctate; following respects: Head: Front as male to punctures confluent, moderately large, shallow, weakly rugulose; setae above eye more pro­ occasionally reduced along midline; setigerous nounced. Clypeal apex slightly narrower and less A REVISION OF THE GENUS Strategus / 141 reflexed than in male. Pronotum: Tubercle virtu­ lection at Glasgow. Type locality: originally ally effaced. Pygidium: In lateral view basal half published as "America" but here restricted to weakly convex. Jamaica.] Scarabaeus simson Fabr., 1787: 5. [Redescrip­ Biology.-Essentially unknown. S. sarpedon tion of Linnaeus' simson.] has been taken on the roots and seed pieces of Scarabaeus titanus Fabr., 1787: 6. [Redescrip­ sugar cane (Stahl and Scaramuzza, 1929; Val­ tion.] des, 1951). Tiphia argentipes Cresson (Hymenoptera: Tiphiidae) and Campsomeris Male.-Length 29.6-40.0 mm; width across tritasciata (Fabr.) (Hymenoptera: Scoliidae) are humerus 14.0-20.0 mm. Color piceous, dull, not two known natural enemies (Valdes, 1951). shining. Head: Front just behind tubercles 'D)stribution.-Cuba. rugose to rugo-punctate to very sparsely punctate; punctures small, shallow. Clypeus with '\"cality Records (Fig. 2).-68 specimens apex slightly reflexed, truncate to subtruncate, examined (27 males, 41 females). Specimens occasionally with a feeble, triangular notch me­ ~ere seen from the following collections: AMNH, dially; surface in majors moderately to densely BMNH, CNCI, HAHC, MCZC, SEMC, USNM. punctate; punctures small or large and confluent C'UBA (67).-CAMAGUEY (25): Baragua, Jaronu, respectively, shallow; surface in minors rugose. Jafi'bonico, Pilar, Santo Tomas Vertientes; Tubercle conical, very low to nearly obsolete, HABANA (6): Cerro, Habana, Santiago de Las transverse, distinctly separated. Mandibles with Vegas, Vedado; LAS VILLAS (22): Cayamas, Cien­ basal lobe small, rounded; middle lobe moderate fuegos; MATANZAS (8): Cardenas; ORIENTE (1): in size, subtriangular; apical lobe small, triangu­ Cristo; No data (5). March (1), April (3), May (36), lar. Interocular width 2.0-2.33 transverse eye June (16), July (3), September (1), October (1). diameters. Mesosternum: Anterior half setiger­ ously punctate. Pronotum: Majors with base Remarks.-The partially punctate mesoster­ lacking a transverse band of sculpturing; entire num, presence of rows of moderate to large pronotum finely granulate, weakly aciculate, punctures on the elytral disc, the rounded sparsely punctate; punctures small, deep; fovea clypeal apex, shining elytra, length of less than deep. Minors as majors except base with a very 30 mm, and absence of horns on the pronotum narrow, punctate band; punctures small to mod­ will serve to separate this species from all others. erately large, sparse, oval to oblong, shallow; Strategus sarpedon is the smallest species in the lateral margin of sides with a similar punctate genus. The males are not dimorphic in the sense band; a punctate to rugo-punctate band extend­ of having major and minor development of pro­ ing postero-medially from anterior angles; fovea notal armature. deep. Horns: Majors (Figs. 79-80) with anterior Although placed in Scaptophilus long, slender, curving forward and slightly to ( = Bothynus) by Chevrolat (1865) and ques­ strongly downward and then upward, apex tioned by others as to its generic placement, S. strongly expanded and forked; horn shaft sub­ sarpedon is indeed a true Strategus. rectangular in cross section, dorsal surface flat to slightly concave and longitudinally carinate on each lateral margin. Posterior horns long, Strategus simson (L.) slender, laterally compressed, attenuate, project­ (Figs. 1,79-82,156-157) ing forward subhorizontally at about 20° from plane of disc; apex acutely rounded; occasion­ Scarabaeus simson L., 1758: 345. [Type material ally a slight ventral flaring just before apex in possibly lost; see remarks. Type locality: origi­ large individuals; in dorsal view horns subparal­ nally published as "America" but actually lel with apices curving toward one another, Jamaica.] bases joined across disc in an arc. Minors (Figs. Scarabaeus manus Fabr., 1775: 10. [Types un­ 81-82) with anterior similar to major except known to me; they are not in the Hunter Col- short, prOjecting forward and upward, apex 142 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

weakly expanded or not, longitudinal carina on ing life history: The eggs are laid singly in spher­ dorsal surface feeble to absent. Posterior horns ical cells of wood fiber formed by the adult in short to subconical, otherwise as majors. Elytra: tunnels made by the males. The average number Sutural stria impressed, crenulate. Disc finely of eggs per female is 15. The average duration of granulate, aciculate, sparsely punctate; incubation is 18 days. Larval development con­ punctures simple and ocellate-umbilicate mixed, sists of three instars: the average length of the the former small and minute mixed, deep, and first instar is 42 days, the second 72 days, and the the latter small to moderate, shallow, occasion­ third 200 days. Larvae are usually found in decay­ allyppsolete; usually 3 feebly impressed striae ing wood and in woody growth at the base of old on lateral half of disc and usually a poorly de­ sugar cane stools where they feed on rotted fined to strong row of ocellate-umbilicate stems and other organic matter. Sugar cane may punctwes mesad of each stria; punctures small suffer considerable damage because of the un­ to moperate, shallow. Sides similar to disc ex­ derground feeding and cutting of cane roots by cep;t;without striae and usually with 2-3 ill­ the larvae; the underground portion of living d~fi;l').ed rows of ocellate-umbilicate punctures as cane stems is usually not attacked. Larvae have on disc. Apex densely punctate; punctures small, also been found in manure. Mature larvae form shallow. Pygidium: Convex in lateral view. Disc pupal cells of wood fiber or soil and undergo a finely granulate, punctate; punctures sparse to pre-pupal stage of from several days to two mod~rate (frequently denser in minors), small, weeks. The average length of pupation is 25 ~.. shallow. Apical margins either side of middle days. After emerging, adults remain in the pupal rugulose in minors, as disc in majors. Genitalia: cells about seven days before burrowing out. Figs. 156-157. Adult food consists of the green parts of trees and occasionally sugar cane. Female.-Length 26.0-37.8 mm; width across In addition to damaging sugar cane (Edwards, humerus 13.8-19.0 mm. As male except in the 1939; Gowdy, 1923), label data indicate that S. following respects: Head: Front rugo-punctate to simson has been taken from banana cargos ar­ rugose. Clypeus with apex narrowly truncate in riving at New York and Washington, D.C. Chapin unworn specimens to narrowly rounded in worn (1932b) suggested that they might possibly be of specimens; surface rugose. Tubercles frequently economic interest in connection with the banana a little larger than in male, and then weakly con­ industry as specimens apparently enter the nected by a feeble, transverse carina. Interocular United States on bunches of the fruit. width 2.33-2.66 transverse eye diameters. Pro­ Gosse (1848), in the only available note on notum: Base with an extremely narrow, punctate natural enemies, found males in the stomachs of to rugo-punctate band; band reduced to basal Nyctibius griseus jamaicensis (Gmelin) (Cap­ bead at middle. Sides just postero-Iateral of rimulgiformes: Nyctibiidae). fovea with a slightly depressed, rounded patch of moderately dense punctures; punctures moder­ Distribution.-Jamaica. Howden (1970) men­ ate to large, oblong to crescent shaped, shallow; tioned that this species was limited largely to lateral margins usually with a wide, punctate to lowland areas. rugo-punctate band in basal half. Anterior third rugose. Fovea shallow. Tubercle very low, trans­ Locality Records (Fig. 1).-68 specimens verse, nearly obsolete. Elytra: Ocellate­ examined (20 males, 48 females). Specimens umbilicate punctures usually less pronounced. were seen from the following collections: AMNH, Pygidium: In lateral view basal 3/4 convex, apical BMNH, CNCI, FMNH, HAHC, MCZC, USNM. V4 concave; protuberant at middle. Disc com­ JAMAICA (68).-CORNWALL (5): Balaclava, Mal­ pletely and moderately densely punctate; verne, Montego Bay, Munro College; MIDDLESEX punctures small to moderate, shallow. Apical (32): Brown's Town, Mandeville, Moneagne, margins rugulose; lateral emargination shallow. Springfields; SURREY (19): Bath, Kingston, Port Biology.-Gowdy (1923), in rather simplified Antonio; No data (12). January (3), March (10), and incomplete observations, noted the follow- May (2), June (5), July (13), August (3), Sep- A REVISION OF THE GENUS Strategus / 143 tember (6), October (7), November (2), December Southern Pines, North Carolina, United (6). States.] Remarks.-$trategus simson is similar to S. caymani, S. aenobarbus, and S. ajax, but it is the Male.-Length 25.0-31.5 mm; width across only species of this group with well-developed humerus 12.0-16.1 mm. Color castaneous, shin­ posterior horns in the majors. Minors may be ing. Head: Front rugo-punctate to rugose, confused with these other species, but the key setigerous above eye. Clypeus with apex slightly characters will separate them. reflexed, moderately to broadly truncate, occa­ The type material for S. simson is apparently sionally with a weak, medial notch; surface fee­ unknown. Chapin (1932b) was apparently of the bly rugulose to rugulose-punctate. Tubercles con­ belief that Linnaeus described S. simson based ical, low to moderate, very transverse, occasion­ I sorely on the works of Browne (1756) and Sloane ally connected by a weak, transverse carina. (1if25) in which the beetle is figured (thus bring­ Mandibles with basal lobe small, rounded; mid­ ing up the interesting possibility that the current dle lobe large, triangular, apex rounded; apical name is invalid because it was based on a lobe small, triangular. Interocular width 3.5-4.0 hypothetical concept, i. e., the illustrations in transverse eye diameters. Mesosternum: An­ Browne and Sloane; since there is no way to terior half setigerously punctate. Pronotum: pl"0ve this, I believe the current name should re­ Base with a narrow rugose band; band reduced main valid). Landin (1956) reported three un­ at middle to basal bead. Disc finely roughened, labeled specimens in the collection of the Lin­ aciculate, sparsely punctate; punctures small to naean Society in London, but there is no way of moderate, shallow to deep. Sides in basal half knowing whether these specimens are the origi­ similar to disc except punctures a little larger; nal types. Although it is possible that these spec­ apical half grading from crescent and oblong­ imens may be the types, the lack of evidence dic­ shaped punctures to rugo-punctate anteriorly; a tates that the types remain unknown. I do not feel Slightly depressed, rounded, rugo-punctate to that a neotype is necessary. rugose area just postero-Iateral of fovea, this Westwood (1837) placed S. simson in junior area occasionally obsolete. Anterior half rugo­ synonymy with S. titanus but in the genus punctate. Fovea shallow. Horns: Anterior a Dynastes; his figured specimens are clearly strong, erect, transverse tubercle, apex usually labeled Jamaica. S. titanus auct. most com­ minutely emarginate. Posterior horns absent. monly referred to S. talpa in the literature up to Elytra: Sutural stria impressed, subcrenulate. the 1930s in reference to specimens from Puerto Disc finely roughened to feebly subgranulate, Rico or the Virgin Islands. aciculate, sparsely punctate; punctures small, shallow. Sides as disc except punctures slightly larger and more dense. Apex subrugose to very densely punctate; punctures small, shallow. Strategus splendens (Beauvais) Pygidium: Convex in lateral view. Disc finely (Figs. 4, 83-84,158-159) granulate,. sparsely punctate; punctures small, shallow to extremely shallow, occasionally with a Scarabaeus splendens Beauvois, 1809: 89. few sparse, moderate-sized pu nctu res. [Types unknown to me. Type locality: South Genitalia: Figs. 158-159. Carolina, United States.] Scarabaeus boscii Beauvois, 1809: 89. [Types Female.-Length 25.2-36.0 mm; width across unknown to me. Type locality: South Carolina, humerus 11.5-18.1 mm. As male except in the United States.] following respects: Head: Front usually more Anastrategus cognatus Casey, 1915: 236. strongly sculptured than in male. Clypeus as in [Holotype female at USNM. Type locality: Kis­ male to apex broadly rounded. Mandibles similar simmee, Florida, United States.] to those of male except middle lobe smaller, api­ Anastrategus carolinensis Casey, 1915: 237. cal lobe usually larger. Pronotum: Tubercle very [Holotype female at USNM. Type locality: low to nearly obsolete. Pygidium: In lateral view 144 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

basal V3 convex, apical 2/3 weakly concave to at the base of the pronotum, length of less than nearly flat. Apical margins either side of middle 35 mm, and lack of posterior horns will easily frequently with sparse, moderate sized separate S. splendens from all other species. punctures. Although Gemminger and Harold (1869) synonymized S. bosci with S. mormon, I believe Biology.-Unknown. Label data indicate spec­ the description of S. bosci clearly indicates that imens have been taken on palms in Florida and it is S. splendens. Arrow (1937a), Casey (1915), injuring young palms in nursery stock. Ritcher and Horn (1875) suspected S. bosci was (1944.~ j 966) has described the larva. synonymous with S. splendens also, but they Distribution.-Southeastern United States. remained undecided. Although S. bosci was de­ scribed prior to S. splendens on the same page Locality Records (Fig. 4).-137 specimens (and more thoroughly as well), it is here formally exam'iJied (56 males, 81 females). Specimens designated that Strategus splendens is the cor­ wer~ ..seen from the following collections: AMNH, rect valid name for this taxon in order to main­ BOR'0, BMNH, CASC, CNCI, FMNH, INHS, KSUC, tain nomenclatural stability. Mcte, OSUC, OSUO, PMNH, SEMC, UMMC, The males do not possess major and minor USNfyl. forms of pronotal development, but some differ­ ences in horn size can be observed. UNlte'1:) STATES (137).-ALABAMA (7): Baldwin " (1): No data; Lee (1): Auburn; Mobile (5): Mobile. " FLORIDA (99): Alachua (2): Gainesville; Brevard Strategus surinamens/s surinamensis (8): Melbourne; Broward (2): Ft. Lauderdale; Burmeister Charlott (2): Punta Gorda; Collier (1): Marco; (Figs. 9, 85-88,160-161) Dade (2): Miami; Duval (3): Jacksonville; Hardee (1): Zolfo Springs; Highlands (5): Archibald BioI. Strategus surinamensis Burmeister, 1847: 135. Stat.; Hillsborough (8): Lutz, Tampa; Holmes (1): [Holotype male at MNHN (Oberthur Collec­ Smyrna; Lee (10): Ft. Myers; Manatee (1): Oneco; tion). Type locality: Guyana.] Okeechobee (1): Okeechobee; Orange (1): Winter Park; Palm Beach (8): Lake Worth, 9 mi. Male.-Length 32.0-40.0 mm; width across W. Lake Worth, West Jupiter; Pinellas (24): Cab­ humerus 16.2-18.0 mm. Color piceous (rarely bage Key, Dunedin, Gulf Port, St. Petersburg; castaneous), feebly shining. Head: Front moder­ . Polk (2): Lakeland; Sarasota (3): Englewood, ately rugose to rugo-punctate, especially behind Sarasota; Volusia (4): Daytona Beach, Enter­ tubercles; setigerous above eye. Clypeus with prize; No data (10). GEORGIA (4): Liberty (1): apex very narrowly truncate, nearly subacute, Hinesville; Macon (1): Oglethorpe; Thomas (1): moderately reflexed; surface weakly rugose to Thomasville; No data (1). MISSISSIPPI (4): Greene rugo-punctate or frequently reduced to sparse (1): Leakesville; Harrison (1): no data; Jackson punctures; punctures small, shallow; when (2): Biloxi. NEW YORK (1): No data. NORTH punctures sparse, surface also finely subgranu­ CAROLINA (16): Moore (16): Southern Pines. late and aciculate. Tubercles conical, low to SOUTH CAROLINA (6): Charleston (2): McClellan­ strong; tubercles when low are transverse, occa­ ville; Georgetown (1): Georgetown; Horry (1): sionally weakly connected by a transverse Myrtle Beach; Richland (1): Columbia; No data carina; tubercles when strong usually not trans­ (1). January (7), February (6), March (23), April verse and are distinctly separated. Mandibles (37), May (15), June (11), July (12), August (3), with basal lobe small, prominently rounded; September (4), November (2). middle lobe large, subtriangular; apical lobe Remarks.-The partially punctate mesoster­ small, triangular. Interocular width 2.0-2.33 num, absence of rows of moderate to large transverse eye diameters. Mesosternum: Com­ punctures on the elytral disc or behind the pletely and setigerously punctate. Pronotum: humerus, the simple and short anterior horn, Base with a narrow to obsolete, rugo-punctate to shining elytra, presence of a narrow rugose band rugose band; band reduced at middle to basal A REVISION OF THE GENUS Strategus / 145

bead. Disc finely subgranulate, aciculate, Sides in basal half with lateral margin rugo­ sparsely punctate; punctures very small, deep. punctate to rugose in a moderately wide band; a Sides as disc to punctures slightly larger. An­ slightly depressed, rounded patch of rugosity terior half as disc in majors; smallest minors not just postero-Iateral of fovea. Anterior half of pro­ seen but presumably with typical increase in notum rugose. Fovea shallow. Tubercle strong, sculpturing. Fovea deep; usually a low, broadly conical, slightly transverse. Elytra: Disc with rounded, longitudinal ridge extending posteri­ punctures slightly larger than in male. Pygidium: orly from base of anterior horn. Horns: Majors In lateral view basal half convex, apical half ;(Figs. 85-86) with anterior long, subslender, at­ weakly to moderately concave. Surface com­ tenuate, curving forward and upward, apex nar­ pletely and setigerously punctate to rugo­ 'rowly rounded; dorsal surface rounded or punctate; punctures small to moderate, very weakly to strongly carinate longitudinally down shallow, moderately dense; setae sparse to "middle. Posterior horns long, stout, laterally moderate, occasionally worn completely off. , .~:compressed, attenuate, apex narrowly rounded; Apical margins rugulose in a narrow band . . ' ,in lateral view horns project upward at about 65° from plane of disc and are frequently recurved Biology.-Unknown. , slightly; in dorsal view horns curve toward one 'another; bases joined across disc in an arc; arc Distribution.-Amazon River north to Ven­ slightly produced at middle. Minors (Figs. 87- ezuela. 88): smallest individuals not seen but presuma­ Locality Records (Fig. 9).-90 specimens bly with typical reduction. Elytra: Sutural stria examined (35 males, 55 females). Specimens impressed, weakly to distinctly crenulate; rarely were seen from the following collections: AHCC, reduced to large, deep, irregularly shaped, AMNH, BCRC, BMNH, CNCI, FMNH, JDG, HAHC, nearly confluent punctures. Disc finely sub­ MCZC, UC, UMMC, USNM. granulate, aciculate, sparsely punctate; punctures minute to very small, shallow; 1-3 very BRAZIL (1 ).-RIO BRANCO (1): Limeo (Rio feebly impressed striae present or not. Sides as SURINAM (1).-No data (1). disc except striae lacking, lateral margin with ECUADOR (1).-NAPO PASTAZA (1): Tena. punctures slightly larger to moderate; 2-3 short January (1). rows of large punctures behind humerus; punctures usually ocellate, simple to umbilicate FRENCH GUIANA (2).-No data (2). (even on same specimen), moderately deep. GUYANA (2).-BERBICE (1): Ituni; ESSEQUIBO (1): Apex densely punctate; punctures small to mod­ Bartica. July (1). erate, shallow. Pygidium: In lateral view basal half convex, apical half weakly convex to nearly SURINAM (1).-No data (1). flat. Disc finely subgranulate, aciculate, sparsely TRINIDAD (37).-CARONI (2): Caparo; MAYARO punctate; punctures minute, shallow, occasion­ (1): Mayaro Beach; ST. GEORGE (28): Arima Val­ ally a few moderate in size. Apical margins either ley, Maracas Bay, Morne Bleu, Orange Grove, side of middle with punctures larger or occa­ Port-of-Spain, St. Augustine; ST. PATRICK (3): sionally weakly rugulose in a narrow band. Fyzabad; No data (3). February (2), March (2), Genitalia: Figs. 160-161. Parameres slightly vari­ April (4), May (3), June (10), August (9), Sep­ able in degree of curvature. tember (1). Female.-Length 31.8-40.7 mm; width across VENEZUELA (46).-ARAGUA (7): Guayas, La Pro­ humerus 14.7-19.0 mm. As male except in the videncia, Maracay, Rancho Grande; DISTRITO following respects: Head: Front coarsely rugose. FEDERAL (11): Caracas, EI Valle; FALCON (15): Clypeus with surface weakly to strongly rugose. Cerroda Mision (EI Alzo), Los Taques, 60 mi. SE. Mandibles similar except middle lobe a little Maracaibo, Palma Sola; MONAGAS (1): Caripito; smaller, apical lobe a little larger. Pronotum: PORTUGUESA (1): Guanare; SUCRE (1): Guanoca; Base with a narrow to moderate, rugose band. TRUJILLO (1): Trujillo; ZUILA (1): no data; No data 146 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

(8). April (2), May (24), June (4), July (5), August Male.-Length 18.5-40.3 mm; width across (1), October (1). humerus 13.0-19.6 mm. Color piceous (rarely Remarks.-This species may be distinguished castaneous), shining. Head: Front strongly by the key characters. Although very similar to S. rugose to rugo-punctate, setigerous above eye. jugurtha, S. surinamensis surinamensis lacks the Clypeus with apex very narrowly truncate, nearly setae on the elytral apices, possesses stouter, subacute, usually feebly emarginate in unworn more erect horns, and is apparently allopatric. specimens; weakly to strongly reflexed; surface Tt'\e genitalia are nearly identical in the two varies from weakly (rarely strongly) rugose to species, but in S. jugurtha the parameres are feebly rugo-punctate to moderately or densely usually slightly more elongate. punctate; punctures moderate in size, very shal­ low. Tubercles conical, moderate to strong, Str~tegus surinamensis surinamensis may be di~ttlJguished from S. surinamensis hirtus by the transverse, usually widely separated. Mandibles with basal lobe small, prominently rounded' foll:cfwing combination of characters: In the males, the base of the pronotum has a narrow to middle lobe large, subtriangular; apial lob~ small, triangular. Interocular width 2.0-2.5 trans­ b6~0lete band of sculpturing in surinamensis and a narrow to moderate band in hirtus' the verse eye diameters. Mesosternum: Completely discal punctures of the pronotum are u~ually and setigerously punctate. Pronotum: Base with small in surinamensis and very small in hirtus; a narrow to moderate rugose to rugo-punctate ... the posterior horns in surinamensis are fre­ band; band reduced to basal bead at middle. ,. quently recurved posteriorly whereas in hirtus Disc finely subgranulate, aciculate, sparsely they are usually not; the discal punctures of the punctate; punctures small, deep. Sides as disc to elytra are small in surinamensis and very small in punctures slightly larger in majors; minors with a hirtus; the pygidium in surinamensis is sparsely moderately wide, rugo-punctate to rugose band and minutely punctate and occasionally with a on lateral margin in basal half; a slightly depres­ few moderate-sized punctures, whereas in hirtus sed, rounded, rugo-punctate to rugose area just the pygidium is sparsely to moderately densely postero-Iateral of fovea in minors, at base of pos­ punctate with the punctures small to moderate in terior horns in majors. Anterior half as disc in size, and there is usually a small median band or majors; minors with a rugo-punctate to rugose triangular patch of large, setigerous punctures band extending postero-medially from anterior on the apical portion on the disc. angles or with anterior half completely rugose. In the females the pygidium of surinamensis is Fovea deep; usually a low, rounded, longitudinal moderately punctate to rugo-punctate whereas ridge extending posteriorly from base of anterior in hirtus the pygidium is densely rugo-punctate horn. Horns: Majors (Figs. 89-90) with anterior to subrugose. long, subslender, attenuate curving forward and I believe that the creation of an Amazonian in­ upward, apex narrowly rounded, dorsal surface land sea during the Pleistocene is probably di­ rounded or weakly to strongly carinate longitud­ rectly responsible for the isolation and sub­ inally down middle. Posterior horns long, stout, sequent subspeciation in S. surinamensis (see laterally compressed, attenuate, apex narrowly section on zoogeography). rounded; in lateral view horns project upwards at about 45-65° from plane of disc, only rarely re­ curved; in dorsal view horns subparallel to curv­ Strategus surinamensis hirtus Sternberg ing toward one another, bases joined across disc (Figs. 9, 89-92,162-163) in an arc, arc slightly produced at middle. Minors (Figs. 91-92) with anterior short, conical, trans­ Strategus hirtus Sternberg, 1910:100. [Holotype verse. Posterior horns low and rounded, or later­ female at ZMHU. Type locality: here desig­ a~ly compressed triangular bosses; in lateral nated as Reyes, Bolivia.] VI~W dorsal surface subhorizontal to sloping Strategus kolbeanus Prell, 1934: 164. [Types un­ slightly forward and downward, anterior edge known to me. Type Locality: Sao Paulo, vertical to subvertical; in dorsal view horns sub­ Brazil.] New synonomy. parallel, bases joined across disc in an arc. A REVISION OF THE: GENUS Strategus / 147

Elytra: Sutural stria impressed, weakly to dis­ AMNH, BCRC, BMNH, CASC, CNCI, FMNH, tinctly crenulate. Disc finely subgranulate, acicu­ HAHC, LACM, MCZC, PMNH, SEMC, UC, UMMC, late, sparsely punctate; punctures small, shal­ USNM, USP, ZMHU. low; 1-3 very feebly impressed striae present or ARGENTINA (19).-CHACO (1): R. S. Pena, COR­ not. Sides as disc except striae lacking, lateral RIENTES (3): Bella Vista; JUJAV (1): Jujay Ciudad; margin with punctures slightly larger to moder­ MISIONES (13): Puerto Bemberg, Rio Alto, ate; 2-3 short rows of large punctures behind Parana; SALTA (1): Salta. January (3), February humerus; punctures usually ocellate, simple to (4), December (12). umbilicate (even on same specimen), moderately deep (4 specimens seen where these punctures BOLIVIA (13).-CHUQUIBACA (2): Yhancaroinza; are nearly obsolete). Apex densely punctate; SANTA CRUZ (10): San Jose de Chiquitos, Santa pu,nctures small to mOderate, shallow. Pygidium: Cruz;·No data (1). In ,~ateral view basal half convex, apical half BRAZIL (138).-AMAZONAS (1): Tefe; DISTRITO Weakly convex to nearly flat. Disc finely sub- FEDERAL (5): Brasilia; GOlAS (7): Dianopolis, . granulate, aciculate, sparsely to moderately den­ Fazenda Nova Orlandia (Jatai), Goiatuba, sely punctate; punctures usually small, occa­ Leopoldo de Bulhoes, Vianopolis; MATO GROSSO sionally moderate, shallow; usually a small, me­ (13): Campo Grande, Corumba, Mato Verde (Rio cHan band or triangular patch of moderately Araguaia), Rio Sao Domingos, Xavantina; PARA lirge, setigerous punctures on apical portion of (1): Cachibbo; PARANA (3): Caviuna, Curitiba, disc, occasionally reduced or obsolete. Apical Rolandia; SANTA CATARINA (71): Nova Teutonia, margins either side of middle weakly rugo­ Rio Natal, Sao Francisco do Sui; SAo PAULO (30): punctate to rugulose. Genitalia: Figs. 162-163. Bilac, Castilho, Divinolandia, Fazenda Cam­ Parameres slightly variable in relative width and pininas (Mogi Guacu), Neves Paulista, degree of curvature. Piracicaba, Ribeirao Preto (Tamandua and FEMALE.-Length 30.0-38.1 mm; width Fazenda Medicina), Sao Paulo; No data (7). across humerus 13.8-18.0 mm. As male except in January (54), February (24), March (2), April (1), the following respects: Head: Front coarsely September (2), October (10), November (13), De­ rugose. Clypeus with surface weakly to strongly cember (23). rugose. Mandibles similar to those of male ex­ PARAGUAY (16).-ALTO PARANA (2): Puerto Ber­ cept middle lobe a little smaller, apical lobe a toni; CAAGUAZU (3): EstanCia Primera, Puerto little larger. Interocular width 2.0-2.66 transverse Central; CONCEPCION (1): Horqueta; GUAIRA (1): eye diameters. Pronotum: Sides in basal half Puerto Suaryro (Villarrica); LA CORDILLERA (2): with lateral margin rugo-punctate to rugose in a Ascuncion, San Bernardino; No data (7). January moderately wide band. Anterior half of pronotum (1), February (1), March (2), May (1), July (1), Oc­ rugose. Fovea moderately deep. Tubercle tober (1), November (1), December (6). strong, conical slightly transverse. Pygidium: In lateral view basal half convex, apical half weakly PERU (5).-HuANUCO (2): Tingo Maria; liMA (1): to moderately concave. Surface completely, M. Sani Beni; LORETO (1): Nauta; SAN MARTIN (1): densely, and setigerously rugo-punctate to sub­ vicinity of Rioja. November (3). rugose; setae dense. Apical margins rugose. Remarks.-This species was commonly Biology.-Unknown. Specimens have been known as Strategus tridens Burm. prior to this taken in coconut nurseries in Brazil (Vayssierre, study, primarily because of Kolbe's (1906) com­ 1965) and at lights. plete description of what he thought Burmeis­ ter's (1847) S. tridens to be. Recourse to the sup­ Distribution.-Amazon River south to Argen­ posed "type" of Burmeister's S. tridens (MNHN) tina. showed it to be S. validus. The "type" of S. tri­ Locality Records (Fig. 9).-191 specimens dens was apparently designated later by some­ examined (68 males, 123 females). Specimens one other than Burmeister and is not actually a were seen from the following collections: AHCC, type at all. Furthermore, the name S. tridens is 148 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM here declared a nomen nudum because Bur­ nently rounded; middle lobe small, broadly meister himself stated that he did not consider rounded at nearly a right angle; apical lobe very Dupont's unpublished manuscript tridens to be small, subtriangular. Interocular width 2.66 any different from S. a/oeus; the name was not transverse eye diameters. Mesosternum: An­ proposed as describing a new taxon by Burmeis­ terior half setigerously punctate. Pronotum: ter but merely mentioned in textual discussion Base with a moderately wide, rugose band, band and does not meet the requirements of Articles reduced at midline to basal bead. Disc very finely 12 and 16 of the 1961 Code. Subsequent authors, subgranulate, aciculate, sparsely punctate; however (Arrow, 1937a; Blackwelder, 1944; punctures small and minute mixed, shallow. Kolbe, 1906), credited Burmeister for validly de­ Sides as disc except lateral margin with a wide, scri'bjng S. tridens although he did not do so. punctate to rugo-punctate band; punctures Ste.niberg's S. hirtus, as the available senior large, shallow, some crescent shaped. Anterior ? syn~>nym, replaces S. tridens as the correct name half as disc. Fovea moderately deep. Horns: An­ forth is taxon. terior long, slender, attenuate, curving forward , ':,think it is interesting to note that Kolbe was and upward, apex narrowly rounded, dorsal sur­ fir,st to accurately describe the taxon S. face rounded. Posterior horns long, subslender, sUfinamensis hirtus, but he was apparently una­ attenuate, laterally compressed, apex rounded; ware of the status of Burmeister's so-called type in lateral view horns project forward and upward and its name, and, therefore, overlooked the op­ at about 60° from plane of disc; in dorsal view portunity of naming the species which he, in fact, horns curve toward one another. E/ytra: Sutural first described. stria impressed, feebly crenulate. Disc weakly Sternberg described the type of S. surinamen­ subgranulate, aciculate, sparsely punctate; sis hirtus as being from Reyes, but he did not punctures small, moderately deep; medial third know whether the Reyes in Mexico or in Bolivia with moderately dense, ocellate-umbilicate was referred to. Reyes, Bolivia, is here desig­ punctures; punctures irregular, large, shallow; nated as the type locality, since S. surinamensis lateral 213 with 6 rows of moderate to large, hirtus is unquestionably South American and not ocellate-umbilicate, shallow punctures. Sides Mexican. with similar background sculpturing as disc and This species may be distinguished from other with 6 confused rows of ocellate-umbilicate species by the key characters and from S. punctures; punctures moderate to large, shal­ surinamensis surinamensis by those characters low, occaSionally confluent. Apex densely listed in the remarks for that subspecies. punctate to rugo-punctate. Pygidium: Convex in lateral view. Disc finely granulate, sparsely punctate medially; punctures minute, small, and Strategus symphenax, new species moderate mixed, shallow; a few moderate, oval (Figs. 2, 93-94,164-165) to oblong, punctures at apex; punctation later­ ally very dense to rugo-punctate. Apical margins Type MateriaJ.-Holotype male, labeled "Las strongly rugo-punctate in a wide band. Genitalia: Martinas, P. de R., Cuba, Jun 24/40, J. Acuna"; Figs. 164-165. deposited at USNM. Also one male paratype de­ posited in BCRC. FemaJe.-Unknown. HoJotype (Figs. 93-94).-Male. Length 35.0 Variation.-Single male paratype as holotype mm; width across humerus 17.0 mm. Color pice­ except in the following respects: Length 37.5 ous, shining. Head: Front densely rugo-punctate mm; width across humerus 17.6 mm. Head: to rugose; a longitudinal, smooth area medially; Setae above eye apparently worn off. Mandibles setigerous above eye. Clypeus with apex broadly with middle lobe obtusely rounded. Interocular truncate, moderately reflexed; surface trans­ width 3.0 transverse eye diameters. Pronotum: versely rugose. Tubercles conical, very low Lateral margin with punctures moderate to large. (nearly effaced), slightly transverse, widely sepa­ Horns: Posterior horns project at about 70° from rated. Mandibles with basal lobe small, promi- plane of disc. E/ytra: Sutural stria strongly crenu- A REVISION OF THE GENUS Strategus I 149

late. Disc on lateral 213 with 6 rows of punctures terior of tubercles usually densely punctate; as in holotype, but rows 3 and 6 confused. punctures small to moderately large, shallow, Pygidium: Disc densely punctate medially. often confluent; front in minors finely rugulose. Clypeus with apex moderately reflexed, strongly Biology.-Unknown. and narrowly bidentate in unworn specimens, Distribution.-Cuba. narrowly truncate or feebly emarginate in worn specimens; surface in majors very finely sub­ Locality Records (Fig. 2).- 2 male specimens granulate, aciculate, sparsely punctate; ~~amined (BCRC, USNM). punctures small, moderately deep; surface in CUBA (1 ).-PINAR DEL RIO (1): Las Martinas; No minors moderately to densely punctate; data (1). June (1) . punctures small to moderately large, shallow, .~ often confluent. Tubercles conical, very low or , ~emarks.-The partially punctate mesoster- virtually effaced, transverse, widely separated. rtum, presence of rows of moderate to large Mandibles with basal lobe small, prominently " Deellate-umbilicate punctures on the elytral disc rounded; middle lobe moderate in size, triangu­ and adjacent to the sutural stria, the broadly lar to subtriangular, apex narrowly rounded; api­ trunctate clypeus, shining elytra, length greater cal lobe small, subtriangular. Interocular width than 30 mm, and pronotum with three simple 2.66-3.0 transverse eye diameters. Mesoster­ h&rns will distinguish this species. num: Anterior half setigerously punctate. Pro­ Although S. symphenax is very similar to S. notum: Base in majors without a rugose band; anachoreta, the form of the horns and body base in minors with a narrow, medially reduced, shape differs, and S. anachoreta is without large punctate band; punctures small to large, shal­ ocellate-umbilicate punctures adjacent to the low, oblong to crescent shaped. Disc in majors sutural stria. In addition, the genitalia of S. very finely granulate, aciculate, very sparsely anachoreta are more stout than those of S. sym­ punctate; punctures minute to small, deep; disc phenax. in minors similar except frequently with an ir­ Etymology.-From the Greek symphenax regular band of sparse punctures extending meaning a partner in deceit; so named because, from base of posterior horns postero-medially to until very late in this study, I believed the speci­ center base of disc; punctures moderate to mens belonged to S. anachoreta, hence the two large, shallow, oval to oblong to crescent species were partners in deceit. shaped. Sides in majors as disc. Sides in minors similar except lateral margin more punctate; punctures large, shallow, crescent shaped; base Strategus syphax (Fabr.) of posterior horn usually with a slightly depres­ (Figs. 3, 104-107, 172-173) sed punctate area; punctures sparse, small to moderately large, crescent shaped. Anterior half Scarabaeus syphax Fabr., 1775: 9. [Holotype in majors as disc; anterior half in minors usually male at BMNH (Banks Collection). Type local­ with a band of rugosity extending postero­ ity: originally published as America but here medially from anterior angle or else variably restricted to Guadeloupe.] rugose. Fovea deep. Horns: Majors (Figs. 104- Scarabaeus syphax Fabr., 1787: 6. [Redescrip­ 105) with anterior very long, slender, attenuate, tion.] curving forward and upward, apex narrowly Scarabaeus vulcanus Fabr., 1792: 11. [Type ap­ rounded, dorsal surface usually rounded (occa­ parently lost. Type locality: Guadeloupe.] sionally almost flat). Posterior horns very long, subslender, attenuate, laterally compressed, Male.-Length 33.6-42.8 mm; width across apex narrowly rounded; in lateral view horns humerus 16.6-21.5 mm. Color piceous, moder­ slightly enlarged ventrally about V3 total distance ately shining. Head: Front in majors very finely from apex and extending forward and upward at subgranulate, aciculate, sparsely punctate; about 35° from plane of disc; in dorsal view punctures small, moderately deep; area just pos- horns curve toward one another. Minors (Figs. 150 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

106-107) with anterior short, slender, attenuate, margins rugo-punctate to rugulose; lateral apex narrowly rounded. Posterior horns reduced emargination shallow. to prominent, triangular bosses; in lateral view Biology.-Unknown. dorsal edge slopes forward and downward, an­ terior edge vertical to subvertical; in dorsal view Distribution.-Guadeloupe. horns laterally compressed, parallel, bases Locality Records (Fig. 3).-10 specimens joined across disc in an arc. Elytra: Sutural stria examined (5 males, 5 females). Specimens were a row of close-set, moderate to large, umbilicate seen from the following collections: BCRC, punctures; punctures not forming an impressed BMNH, MNHN. F. Chalumeau (personal com­ line. Disc finely granulate, aciculate, sparsely to munication) provided 14 additional records. moderately punctate; punctures minute to small, shallbw, mixed with moderate to large, shallow, GUADELOUPE (24).-GRANDE TERRE (2): Grands oC!=lltate-umbilicate punctures; some ocellate­ Fonds, Pointe-a- Pitre (Chalumeau, in !itt., and umbilicate punctures in distinct rows. Sides simi­ Paulian, 1947, respectively); GUADELOUPE (15): larexcept ocellate-umbilicate punctures greatly Goyave (Chalumeau, in !itt.), IFAC (Chalumeau, reduced in number. Apex densely punctate; in litt.), Petit-Bourg, Sainte Rose, Sofa'ia pLinctures small to moderately large, shallow, (Chalumeau, in litt.), Trois Rivieres (Paulian, usually confluent. Pygidium: In lateral view basal 1947), Vernou (Chalumeau, in litt.); No data (7). half convex, apical half nearly flat to feebly con­ April (1), May (1), June (7), July (1), September cave. Disc aciculate, finely granulate, (1), October (1). roughened, moderately densely punctate; Remarks.-The bidentate clypeal apex, punctures minute to small, very shallow; majors punctate sutural "stria," and large discal frequently with a subapical, median, longitudi­ punctures on the median third of the elytral disc nal, shallow depression. Apical margins either are diagnostic for this species. side of midline rugose; lateral emargination shal­ Verill's (1906) S. tricornis was questionably low. Genitalia: Figs. 172-173. synonymized with S. vulcan us by Arrow (1911, 1937a) and positively by Leng and Mutchler Female.-Length 32.0-40.5 mm; width across (1917), but it is here removed from synonymy; it humerus 15.5-20.3 mm. As male except in the is definitely not S. syphax. following respects: Head: Front punctate to Fleutiaux and Salle (1889) may have applied rugo-punctate; punctures moderate to dense, the name S. syphax to S. vulcan us although their small to large, shallow, often confluent. Clypeus checklist differentiates between syphax and with surface as front. Mandibles similar to those as. vulcanus. Lacordaire (1856) considered S. of male except middle lobe slightly smaller. Pro­ as. vulcan us a variety of S. syphax thus accurately notum: Base with a narrow, punctate to rugo­ placing S. vulcanus in synonymy with S. syphax. punctate band; punctures sparse to dense, small Illustrations of the type of S. syphax were not to large, oval to oblong. Disc sparsely punctate; seen until very late in this study and after the punctures small, deep; an irregular band of plates had been prepared, hence the interruption moderately dense punctures extending from lat­ in alphabetical sequence in the habitus figures. eral base of fovea postero-medially to center base of disc; punctures moderate to large, shal­ low, oval to oblong. Sides in basal half punctate to rugo-punctate; punctures moderate in den­ Strategus talpa (Fabr.) sity, small to large, shallow, frequently crescent (Figs. 3,95-97,166-167) shaped; a slightly depressed, rounded patch of rugosity postero-Iateral of fovea. Anterior half Scarabaeus talpa Fabr., 1792: 32. [Lectotype rugo-punctate. Fovea shallow. Tubercle conical, male, labeled "talpa," and lectoallotype transverse, strong. Pygidium: Convex in lateral female, labeled "talpa," here designated; de­ view. Disc moderately densely punctate; posited at UZM with my lectotype labels. Type punctures small to moderate, shallow. Apical locality: St. Barthelemy (West Indies).] A REVISION OF THE GENUS Strategus I 151

Strategus barbigerus Chapin, 1932b; 455. horns are large, pyramidal bosses; in lateral view [Holotype male and 24 paratypes at USNM; 20 dorsal surface slopes downward and backward, paratypes at AMNH. Type locality: Aguirre, anterior surface subvertical; in dorsal view horns Puerto Rico.] New synonymy. parallel, laterally compressed, bases jOined across disc in an arc. Minors (Fig. 97) with an­ Male.-Length 27.0-37.0 mm; width across terior short to tuberculate; when short the an­ humerus 13.3-19.0 mm. Color piceous, dull, not terior is stout, sides subparallel, apex bifid but shining. Head: Front of majors sparsely not expanded, dorsal surface rounded. Posterior punctate; punctures small to moderate, shallow, horns vary from low, pyramidal bosses to com­ becoming denser and larger just behind tuber­ pletely absent; when low, bosses are similar to cles, setigerous above eye in unworn specimens; majors except much reduced. Elytra: Sutural minors with front coarsely punctate to rugo­ stria impressed, crenulate. Disc very finely punctate, also setigerous above eye in unworn granulate, aciculate, sparsely punctate (rarely :. specimens. Clypeus with apex narrowly subtrun­ moderately dense); punctures simple and um­ cate to truncate, weakly reflexed; surface in bilicate mixed, small (rarely small and minute majors moderately to moderately densely mixed), shallow; disc on lateral 2/3 with 6 rows of punctate; punctures moderate to large, shallow; moderate to large, umbilicate, shallow surface in minors rugo-punctate to rugose. punctures, rows 1-2 and 4-5 regularly punctate >"Tubercles conical, very low, slightly transverse, and closely adjacent, rows 3 and 6 irregular and widely separated. Mandibles with basal lobe more sparsely punctate; rows of punctures and small, rounded; middle lobe moderate in size, size of larger punctures occasionally reduced so subtriangular, apex narrowly rounded; apical that rows become confused and difficult to de­ lobe very small, triangular. Interocular width 3.0 fine. Sides similar to disc except row 3 usualy transverse eye diameters in minors grading to reduced to only a few irregular punctures, row 6 4.0 transverse eye diameters in majors. Mesos­ sparsely and irregularly punctate. Apex sparsely ternum: Anterior 1/2_3/4 setigerously punctate. punctate; punctures small, shallow. Pygidium: Pronotum: Base with a very narrow to nearly ob­ Convex in lateral view. Disc finely subgranulate solete rugo-punctate band; band reduced at to granulate, sparsely punctate (especially in middle to basal bead. Disc aciculate, punctate; majors); punctures small to moderate, very shal­ punctures moderate in density, small, shallow; low to effaced. Genitalia: Figs. 166-167. Para­ occasionally a few moderate to large punctures mere just before apex mayor may not be slightly along midline. Sides as disc except majors with a flared laterally. few moderate to large, crescent-shaped punctures at base of posterior horns; minors Female.-Length 25.8-38.0 mm; width across with a slightly depressed, rounded, punctate to humerus 12.0-21.0 mm. As male except in the rugose patch just postero-Iateral of fovea; lateral following respects: Head: Front rugose. Clypeus margin in majors with a band of sparse with apex rounded; surface rugose. Tubercles punctures; punctures moderate to large, simple conical, low, strongly transverse, connected by a to oblong, shallow; lateral margin in minors var­ weak transverse carina. Mandibles similar to ies from densely punctate to rugo-punctate to those of male but middle lobe smaller. Interocu­ rugose. Anterior third as disc in majors; minors lar width 2.66-3.0 transverse eye diameters. Pro­ with a postero-medial, rugo-punctate to rugose notum: Sides with lateral margins punctate to band extending from anterior angles or else with rugo-punctate to rugose in a wide band or sides anterior third of pronotum rugose. Fovea deep in completely punctate to rugo-punctate; majors, moderate to very shallow in minors. punctures moderate to large, moderately dense, Horns: Majors (Figs. 95-96) with anterior long, shallow. Anterior third rugose. Fovea shallow to stout, dorso-ventrally compressed, apex strongly nearly obsolete. Tubercle conical, very low, expanded and bifid, curving forward and up­ transverse. Elytra: Umbilicate punctures fre­ ward; dorsal surface flat, lateral margins de­ quently less pronounced. Pygidium: In lateral lineated by a small, longitudinal carina. Posterior view basal 3/4 convex, apical Y4 slightly concave; 152 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

protuberant at middle. Disc aciculate, very finely onstrated a preference for feeding on rotting subgranulate, sparsely to moderately densely wood and were most commonly found in decay­ punctate; punctures small to large, shallow, fre­ ing tree stumps. Unlike S. oblong us with which it quently elongate in lateral emarginations. Apical is sympatric, larvae of S. talpa can survive on margins rugo-punctate to rugose with a few manure and are frequently found there. The short setae medially in unworn specimens; lat­ grubs also show a preference for the rotten eral emargination very shallow. stems in the stool of sugar cane; they do not usually attack living stems. However, the under­ Biology.-Smythe (1920) conducted rearing ground portions of living stems will be attacked experli'nents in Puerto Rico and established the if the available dead stems are entirely con­ following life history: The eggs are pearly white, sumed by the developing larvae before they opaque, and oblong-oval in shape. When first reach maturity. Larvae have been observed to deposited they averaged 2.8-3.0 x 3.5-3.8 mm; sever completely a cane stalk underground and during< tncubation eggs expanded and just prior then to tunnel upward within the stalk a distance to ha'tthing averaged 4.0-4.5 x 5.3-5.5 mm. The of several inches. Pupae have been found within duraHon of incubation in 207 eggs was 15-19 the excavated underground base of the stalk. days (average 17.5 days). The eggs are laid singly A pupal cell is usually constructed just prior to in spherical cells of soil or fiber which are 2-3 pupation near where the grub stops feeding. The times ,larger than the newly deposited egg. Pre- pupal cell is oblong and somewhat larger than , ferred deposition sites for the eggs are in the the pupa. A quiescent pre-pupal stage lasts from , " chewed-up fiber in which the adult has been several days to several weeks, and during this tunneling or in the excavated interior of buried stage the pre-pupa is very susceptable to attack sugar cane stalks. by fung'i, bacteria, mites, and nematodes. The Larvae are about 8 mm in length when they pupal period ranged from 22 to 29 days (average hatch. Tables 3 and 4 indicate larval develop­ 24 days). Newly emerged adults became active ment. within a week after eclosion and then im­ mediately dug their way out of their chamber. TABLE 3 Adult female longevity in 11 individuals aver­ INSTAR DURATION IN DAYS FOR S. talpa aged 37.5 days (maximum 93 days). The total life span averages about one year. The pre­ INSTAR NO. OBSERVED RANGE AVERAGE oviposition period for five females ranged from 1 115 24-72 40.5 20 to 27 days (average 24 days). The average 2 67 43-85 72.0 number of eggs laid per female was 13, the 3 55 137-282 199.0 maximum number 43. The oviposition period lasted 1-21 days (average eight days). TABLE 4 The adults feed largely on the green parts of INSTAR MEASUREMENTS IN MM FOR S. talpa shrubs and trees but not on foliage. They rarely damage sugar cane, and then only incidentally HEAD CAPSULE W,DTH INSTAR No OBSERVED MAXIMUM BODY LENGTH as they penetrate the cane to deposit eggs or RANGE AVERAGE occasionally to feed on underground stems. 20 3.25-3.90 3.54 25.0 Natural enemies are few in number (see biol­ 2 91 5.642-6.243 45.0 ogy for S. oblongus). The green muscardine fun­ 3 gus, Metarrhizium anisopliae (Metschn.), is par­ 2Less than one week in age. ticularly virulent and attacks the adults as well as 30ver nine weeks in age. the immature stages. The pupa appears to be the most susceptable to attack, and the egg the least The first instar is apparently able to subsist with susceptable. Larvae in the laboratory frequently no more organic materal present than the humus became diseased and died due to the bacterium in black soil although it begins to feed on rotten Micrococcus nigrofasciens which caused shin­ wood near the end of the stadium. Larvae dem- ing, black, hardened areas to form on the body, A REVISION OF THE GENUS Strategus / 153

head, and legs. Campsomeris atrata (Fabr.) Dlstrlbutlon.-Puerto Rico, Virgin Islands, St. (Hymenoptera: Scoliidae) is parasitic on the lar­ Barthelemy, Antigua, and probably other islands vae of S. talpa (Wolcott, 1936, 1948). An unde­ in the Leeward Islands. termined species of predatory mite was observed Locality Records (Fig. 3).-98 specimens to feed on the adults; another undetermined examined (41 males, 57 females). Specimens species was seen to feed on the eggs of the bee­ were seen from the following collections: tle resulting in the death of the egg. Other pred­ AMMH, BCRC, BMNH, CASC, CNCI, HAHC, ators include the Puerto Rican grackle, Quis­ MCZC, USNM, UZM. calus niger brachypterus (Cassin), which feeds on the larvae; the mongoose, Herpestes bir­ LEEWARD ISLANDS (16).-ANTIGUA (1); ST. manicus (Thomas), which feeds on the adults; BARTHELEMY (2); ST. CROIX (7): Christiansted, and the American Giant Toad, Buto marinus (L.), Const. Hill, no data (Beatty, 1944); ST. JOHN (1); which also feeds on the adults. In a detailed ST. THOMAS (3); TORTOLA (2). May (1), June (2), ~tl,l{jy in Puerto Rico, Dexter (1932) found seven August (3), October (3). ,St'rategus (probably S. talpa) in 301 toads for a PUERTO RICO (82).-AGUADILLA (3): Aguadilla, . total of 0.6% of the total bulk found in all toad Isabel a, Las Marias; ARECIBO (2): Dorado, Vega s~mples. Wolcott (1948, 1950) noted that Buto Baja; GUAYAMA (11): Aguirre, Aibonito, marinus was a very effective predator of S. talpa Caguas; HUMACAO (2): EI Verde Camp (Rio in. Puerto Rico and that the beetles nearly disap­ Grande) (Wolcott, 1941), Fajardo (Wolcott, peared when the toads were numerous and vice 1936); ISLA MONA (3): Sardinera (Ramos, 1946); versa. Miskimen and Bond (1970) indicated that MAYAGUEZ (17): Guanica, Mayaguez, Route S. talpa did not reach economic levels of impor­ 334 (Guanica Forest), San German; PONCE tance on St. Croix because of the toad. On the (16): Como Springs, 5 mi. NE. Jayuya, Ponce, other hand, Simmonds (1969) stated that there Santa Isabel; SAN JUAN (7): Bayamon, Corozal, was no definite evidence as to the degree of re­ Rio Piedras, San Juan; VIEQUES ISLAND (2) duction of any specific pest by the toad. (Wolcott, 1923, 1936); No data (19). January Strategus talpa is known as the sugar cane (1), February (3), March (1), April (7), May (11), rhinoceros beetle in Puerto Rico. It is not, and June (9), August (5), September (3), October probably never has been, a serious pest of sugar (14), November (4), December (6). cane even though the larvae were previously found in cane fields. As Smythe (1920) pOinted Remarks.-The partially punctate mesoster- out, the larvae of the rhinoceros beetle were num, rows of moderate to large umbilicate often blamed for the damage caused by Phyl­ punctures on the elytral disc, narrowly sUbtrun­ lophage spp. (Scarabaeidae) which were more cate to truncate clypeal apex, dull elytra, ab­ numerous (but not so readily found) than the lar­ sence of a well-developed angular prominence vae of Strategus. Strategus larvae were found on the lateral margin near the apex of the male among the underground stems of sugar cane genitalia, and the distinctly expanded and forked because of the large amount of organic food ma­ apex of the anterior horn will serve to separate terial that occurs there; the larvae do not feed on this species from all others. cane roots, and damage that does occur is Chapin (1932b) described S. barbigerus with­ caused largely by inadvertent severing of the out seeing the type of S. talpa. roots during feeding. Wolcott (1948) noted that "Strataegus (sic) grubs primarily feed on rotten wood, especially the roots and stumps of trees in Strategus tarqulnius, new species the soil, but when these have been consumed in (Figs. 3, 98-99, 168-169) recently cleared land, readily attack old cane stalks and decaying cane seed, and mayeventu­ Type Material.-Holotype male, labeled ally attack live cane root-stalks and rootlets "Granada (sic), May, 1918"; deposited at AMNH. when other more acceptable sources of organic Allotype female, labeled "Chantilly Est. (Wind­ '11atter are lacking." ward side), Grenada, W. I., H. H. Smith, 60" and 154 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

"W. Indies, 1900.40"; deposited at BMNH. Also teriorly. Fovea shallow. Tubercle conical, low, one male paratype (Figs. 98-99) and one female transverse. Pygidium: In lateral view basal half paratype deposited in BCRC. weakly convex, apical half nearly flat. Disc very finely subgranulate, virtually impunctate me­ Holotype.-Male. Length 32.6 mm; width dially, sparsely punctate on apical margins and across humerus 16.2 mm. Color castaneous, lateral emargination; punctures moderate in shining. Head: Front very feebly rugose behind size, shallow; lateral emargination shallow. tubercles; setigerous above eye. Clypeus with apex very narrowly truncate, moderately re­ Variation.-Males (1 paratype): Length 33.7 flexed; surface sparsely punctate; punctures mm; width across humerus 16.5 mm. Head: Mid­ small, shallow. Tubercles low, transverse, dis­ dle lobe of mandible subtriangular, apex nar­ tin,ctly separated. Mandibles with basal lobe rowly rounded. lnterocular width 3.0 transverse sm~all, prominently rounded; middle lobe moder­ eye diameters. Pronotum: Disc not aciculate. ~e in size, triangular, apex rounded; apical lobe Horns: Anterior curves forward and upward (not .. small, subtriangular. lnterocular width 2.33 downward). Posterior horns in lateral view ex­ transverse eye diameters. Mesosternum: Com­ tend forward and upward up about 35° from pletely and setigerously punctate. Pronotum: plane of disc; in dorsal view horns curve toward Base without sculpturing. Disc very finely sub­ one another. Elytra: Disc with 1 feebly impressed granulate, aciculate, very sparsely punctate; stria. punctures very small, shallow. Sides and anterior Females (1 paratype): L~ngth 26.9 mm; width half as disc. Fovea deep. Horns: Anterior very across humerus 11.8 mm. Head: lnterocular long, slender, attenuate, curving forward and width 2.0 transverse eye diameters. Pronotum: downward and then upward, apex narrowly Base with a narrow, feeble, punctate band; rounded, dorsal surface rounded. Posterior punctures moderate in size, shallow. Sides with horns very long, slender, attenuate, laterally marginal punctate band narrow; punctures compressed, apex narrowly rounded, projecting moderate in size, shallow. Pygidium: Disc mod­ forward and upward at about 20° from plane of erately punctate; punctures small to moderate, disc; in dorsal view horns diverge, bases joined shallow. across disc in an arc. Elytra: Sutural stria Biology.-Unknown. Found in rotting wood impressed, subcrenulate. Disc very finely sub­ (Arrow, 1900). granulate, very sparsely punctate; punctures minute, shallow; lateral half with 3 very feebly Distribution.-Grenada (West Indies). impressed, incomplete striae. Sides similar ex­ Locality Records (Fig. 3).-4 specimens cept punctures slightly larger and striae absent. examined (2 males, 2 females). Specimens de­ Apex moderately punctate; punctures small, posited in the following collections: AMNH, shallow. Pygidium: Convex in lateral view. Disc BCRC, BMNH. subgranulate, very sparsely punctate; punctures minute, shallow. Genitalia: Figs. 168-169. GRENADA (4).-Chantilly Est. (Windward side) (1), Grenville (Windward side) (1), No data (2). Allotype.-Female. Length 33.3 mm; width May (1). across humerus 15.4 mm. As holotype except in the following respects: Head: Front feebly Remarks.-The completely punctate mesos­ rugose. Mandibles similar to those of holotype ternum, absence of rows of punctures on the except middle lobe smaller. Pronotum: Base disc of the elytra and behind the humerus, nar­ with a very narrow band of sparse punctures and rowly truncate clypeal apex, and the lack of rugae; band reduced at middle to basal bead. sculpturing at the base of the pronotum will eas­ Sides in basal half as disc except margin with a ily separate S. tarquinius from all other species. feeble, narrow, punctate band; a rounded, It has the least amount of sculpturing of any slightly depressed, rugo-punctate patch just species in the genus as well as being lighter in postero-lateral of fovea. Anterior half of pro­ color. It seems to be closely related to S. notum rugo-punctate grading to rugose an- jugurtha. A REVISION OF THE GENUS Strategus / 155

Arrow (1900), in referring to the two female de­ terior half setigerously punctate. Pronotum: scribed here, speculated that they were S. fas­ Base without sculpturing (primarily in majors) or cinus or a closely related species and that males with a very narrow, rugo-punctate to rugose were needed to place them definitely; he was band; band reduced at middle to basal bead. apparently unaware that he was dealing with a Disc acciculate, sparsely punctate; punctures new species. small, deep. Sides in majors as disc except The male paratype illustrated here (Figs. punctures slightly denser and larger along mar­ 98-99) was reared from a larva collected by John gin; sides in minors with lateral margin punctate Glaser who first provided me with an example of to rugo-punctate in a moderate to very wide this new species. band in basal 2/3; punctures moderate to dense, small to large, shallow to deep, simple to cres­ Etymology.-From the Latin tarquinius mean­ cent shaped; a rounded, slightly depressed ing proud; here named in reference to the mag­ rugo-punctate to rugose area at base of poste­ 'nificent set of horns on the males. rior horn. Anterior half of pronotum as disc in majors; minors with anterior third variably Strategus validus (Fabr.) rugose. Fovea moderate in depth. Horns: Majors (Figs. 8, 100-103, 170-171) (Figs. 100-101) with anterior very long, slender, , attenuate, curving forward and upward, apex Scarabaeus validus Fabr., 1775: 6. [Holotype narrowly rounded; dorsal surface usually with 1 female at BMNH (Banks Collection). Type loc­ median and 2 lateral parallel, longitudinal, feeble ality: Brazil]. carinae. Posterior horns long, subslender to st­ Scarabaeus tricornis Jablonsky (not Herbst), out, attenuate, laterally compressed, apex 1785: 269. [Types unknown to me. Type local­ rounded; in lateral view horns extend forward ity: unknown, but here presumed to be east­ and upward at about 70-90° from plane of disc, central South America.] occasionally recurving slightly; in dorsal view Scarabaeus validus Fabr., 1787: 4. [Redescrip­ horns slightly diverging to curving toward one tion.] another. Minors (Figs. 102-103) with anterior Oryctes faunus Billberg, 1820: 383. [Types un­ short to conical, suberect, apex acutely rounded, known to me. Type locality: originally pub­ dorsal surface rounded. Posterior horns very lished as Barbary (erroneous), but here pre­ short and triangular or weak to strong triangular sumed to be east-central South America.] bosses or obsolete; in lateral view dorsal edge Strategus tridens Burmeister, 1847: 133. [Nomen horizontal to sloping forward and downward or nudum. See remarks for S. surinamensis hir­ sloping backward and downward, anterior edge tus.] vertical to sloping upward and backward; in dor­ sal view horns or bosses laterally compressed, Male.-Length 31,,0-49.6 mm; width across subparallel, bases joined across disc in an arc. humerus 15.0-24.3 mm. Color piceous, shining. Elytra: Sutural stria strongly impressed, crenu­ Head: Front rugo-punctate to rugulose, sculptur­ late. Disc aciculate, frequently feebly and finely ing usually reduced at midline. Clypeus with subgranulate, sparsely punctate; punctures apex broadly truncate, occasionally very weakly small to moderate and minute mixed, shallow to notched medially, strongly reflexed; surface in deep; lateral half of disc usually with 1-3 very majors moderately punctate; punctures small, feebly impressed, incomplete striae; occasion­ shallow, or surface rugo-punctate; punctures ally a short longitudinal row of small to large large, shallow; surface in minors usually feebly punctures at base just medial of humerus. Sides rugulose. Tubercles conical, strong, widely similar to disc except 1-5 short, confused rows separated. Mandibles with basal lobe small, of punctures behind humerus; punctures usually prominently rounded; middle lobe large, sub­ sparse, moderate to large, frequently ocellate­ triangular, apex narrowly rounded; apical lobe umbilicate, moderately deep. Apex moderately small, triangular. Interocular width 2.0-2.25 densely punctate; punctures small, shallow. transverse eye diameters. Mesosternum: An- Pygidium: Convex in lateral view. Surface acicu- 156 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM late, finely subgranulate, sparsely punctate; AMNH, BCRC, BMNH, CASC, CNCI, FMNH, punctures minute to small, shallow. Genitalia: HAHC, LACM, MCZC, PMNH, UC, UMMC, URU, Figs. 170-171. Parameres vary from subslender USNM,USP. . to stout and from subparallel to slightly arcuate. ARGENTINA (3).-BuENOS AIRES (1): Buenos Female.-Length 32.0-47.0 mm; width across Aires; JUJUY (Hayward, 1942); MISIONES (1): humerus 15.0-25.0 mm. As male except in the Puerto Aguirre; RIo NEGRO (1): Negro Muerto; following respects: Head: Front coarsely rugo­ SALTA (Hayward, 1942); TUCUMAN (Hayward, punctate to rugose. Clypeus with apex broadly 1942). November (1), December (1). sUbtruncate to truncate, but not so broadly as in BRAZIL (348).-AMAZONAS (6): Ipiranga; CEARA male; surface coarsely rugo-punctate to (Goncalves, 1946); ESPIRITO SANTO (3): Cor­ ruguJose. Mandibles similar to those of male ~x­ rego do Ita, Fazenda Corrego Alto (C. Itape cept(middle lobe smaller, usually subequal with Mirim); GOlAS (1): Goias; MATO GROSSO (3): apreal lobe. Interocular width 2.~2.5 transverse Albuquerque, Guape, Km. 100 (Br. 55); MINAS ~ye diameters. Pronotum: Base with a narrow to GERIAS (4): Belo Horizonte, Vicosa; PARA (2): moderate rugo-punctate to rugose band; band Belem, Fazenda Santa Maria (Monte Alegre); reduced at middle to basal bead. Sides in basal PARANA (37): Banhados, Caviuna, Curitiba, 1/2-2/3 moderately to densely punctate; Ponte Grossa, R.R. from Curitiba to Parangua, pu~ctures moderate to large, shallow, frequently Rolandia; PIAUI (Goncalves, 1946); RIO DE crescent shaped; a slightly depressed, rounded, JANEIRO (2): Sarada Carioca; RIo GRANDE DO rugo-punctate to rugose area postero-latera.1 of SUL (17): Pelotas, Rio Grande, S. Lorenzo; fovea, lateral margin with a moderate to wide, SANTA CATARINA (130): Corupa, Joinville, Nova rugo-punctate to rugose band in basal half. An­ Teutonia, Pinhal, Rio Verhelo; SAo PAULO terior 1/3-1/2 rugose. Fovea shallow. Tubercle (124): Alto de Serra, Barveri, Campininas conical, low, transverse. Elytra: Disc as in male (Mogi Guacu), Cantareira, Caraguatatuba, to lateral half more strongly punctate with 1-5 Cipo, Divinolandia, Est. BioI. Borac~a incomplete, confused rows of punctures; (Salesopolis), Fazenda Medicina (Ribeirao punctures moderate to large, simple to PretO), Fazenda Paul D'Alho (Itu), Itu, Pi_n­ ocellate-umbilicate, deep. Sides behind humerus damonhangaba, Piracicaba, Rio Claro, Sao with 3-6 incomplete, confused rows of similar Jose dos Campos, Sao Paulo, Serra Negre; No punctures. Pygidium: Weakly convex in lateral data (19). January (62), February (50), March view. Disc aciculate, finely subgranulate, (9), April (4), June (1), July (1), September (11), sparsely to moderately punctate; punctures October (4), November (30), December (99). moderate to large, simple to oval to oblong, shal­ low. Lateral emargination very shallow. PARAGUAY (1).-CAAGUAZU (1): Estancia Prim­ era. January (1). Biology.-Strategus validus has been taken from the base of wax palms in Brazil (Goncalves, URUGUAY (10).-ARTIGAS (1): Rio Cuareim; 1946) and from sugar cane, bananas, pineapples, MALDONADO (3): Maldonado; RIVERA (3): Ar­ and coconut palms in Argentina and Brazil royo de la Aurora (Sierra de la Aurora), Camino (Costa Lima, 1953; Hayward, 1942). Costa Lima a Tranqueras, Rivera; TACURAREMBO (3): Pun­ also indicated that this species is an inter­ tas Arroyo Laureles. mediate host for Macracanthorhynchus hirudinaceus (Pallas) (Archiacanthocephala: Remarks.-The key characters will distinguish Oligacanthorhynchidae). Strategus validus from all other species although the erect horns, general absence of basal Distribution.-Argentina, Brazil, Paraguay, sculpturing on the pronotum, and the broadly Uruguay. truncate clypeus will usually be sufficient to Locality Records (Fig. 8).-362 specimens separate this species. S. validus is closp,ly related examined (155 males, 207 females). Specimens to S. mandibularis, but the greater degree of were seen from the following collections: AHCC, basal sculpturing on the pronotum and the much A REVISION OF THE GENUS Strategus / 157

larger middle lobe of the mandibles of the latter other species that S. verrilli might be confused will easily separate the two. with, and so I feel confident in renaming the Based on the description only, Arrow (1914) taxon now during this revision rather than wait­ was probably correct in synonymizing Oryctes ing until additional specimens are taken. I be­ faunus Billberg with S. validus. lieve, based on the description and photographs, that this species is valid and may be closely re­ Strategus verrilli, new name lated to S. jugurtha or S. tarquinius. The lack of (Fig. 3) more specific knowledge regarding certain character states prevents it from being included Strategus verrilli, New name. [Replaces in the key to the species. The precipitous terrain , Strategus tricornis (Verrill), 1906: 317, which is of Dominica has undoubtedly aided in delaying , a :secondary junior homonym of Strategus the rediscovery of S. verrilli, and even the 't;icornis (Jablonsky), 1785: 269, which is, in Bredin-Archibald-Smithsonian Biological Survey , ··"~urn, a junior subjective synonym of Strategus of Dominica failed to yield any specimens. '''yalidus (Fabr.), 1775. Type locality: Dominica • (West Indies).] Etymology.-The species is renamed after A. Hyatt Verrill who originally described the species. i~emarks.-Arrow (1911, 1937a) questionably synonymized Verrill's new Dominican species ZOOGEOGRAPHY with S. vulcan us ( = syphax) (see remarks for S. syphax). All of Verrill's type material has been This section will attempt to sketch briefly the lost or destroyed (Kirby Brown, personal com­ distribution of the species of Strategus primarily munication), and apparently no male specimens within the framework of an historical analysis resembling this species have been taken from where geology and chorology are the principal Dominica since. I do have one female specimen parameters. The model proposed here is based from Dominica which does not really seem to fit largely on logical inferences drawing upon what any other species, but I currently have no reliable I believe to be sufficient circumstantial evidence; way of placing it in S. verrilli. Verill indicated that it may serve for testing the distributional patterns the female was unknown when he described the of other groups. species in 1906. This specimen shall be retained Wickam's (1914) single Miocene (actually until additional specimens are taken. Strategus Oligiocene) fossil from Colorado places the verrilli is definitely not Strategus syphax of genus at least 12-25 million years B.P., but if the neighboring Guadeloupe because Verrill's origi­ fossil is indeed a true Strategus (which it appears nal description (1906) and subsequent photo­ to be), then the age of the genus is certainly graphs (1907) clearly show very smooth, shining older. elytra as opposed to the sculptured elytra of S. In the absence of any hard evidence, it is here syphax. S. verrilli is also not S. tarquinius for the suggested that Mexico-Central America is the following reasons: the body shape of S. tar­ place of origin for the genus because this area quinius is proportionately more oblong; the pos­ contains a modest number (four) of primitive terior horns of S. tarquinius are considerably species whereas all other areas (excluding the more erect; the body and legs of S. tarquinius Antilles) contain more of the derived species (see are light reddish brown and S. verrilli is a deep Table 6). Kolbe (1906) also proposed the north­ black; the head tubercles ofS. tarquinius are dis­ ern subtropics as the area of origin for the genus tinctly separated whereas they form a transverse because most of the hornless (primitive) species ridge in S. verrilli; the measurements given for S. occurred there. The Antilles are ruled out as a verrilli indicate that it is at least 10-20% larger center of origin for the genus despite the high than S. tarquinius. number (eight) of primitive species occurring Other than S. syphax and S. tarquinius and there. The Greater Antilles were not even emer­ perhaps S. jugurtha (which has easily distin­ gent until the early Miocene (Woodring, 1954), guishable post-humeral punctures), there are no and it would therefore be impossible to have an 158 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

earlier Oligocene fossil in Colorado as is the isolated in the more humid mountain areas. case with S. cessatus. I believe the majority of Their ecological valency has undoubtedly al­ more primitive Caribbean species is indicative of lowed for adaptation to the cooler temperatures a longer and continued isolation as will be men­ of the slightly higher elevations (especially dur­ tioned later. Minor supporting evidence for a ing the Pleistocene glaciations), but as Dillon Mexican-Central American origin is the fact that (1956) and Howden (1963, 1966) have pointed this area.-,is about the center of the range for the out, the desert areas had by this time formed genus, thus allowing for radial expansion; in ad­ impassable barriers to further dispersal. dition, the second largest number of species oc­ On the other hand, Martin and Mehringer curs in thi,s'region which tends to indicate that (1965), using extensive and more recent data on the genu~ f1Iay have been here longer than in any Southwestern pollen profiles, have decisively other ~iea (the larger number of endemic shown a direct pollen correlation with the Wis­ specie~ ;in the Antilles is a misleading number consin glaciation, starting about 70,000 years clue and has been explained by greater isolation B.P., which indicates a 900-1,200 m lowering of .' due to fragmented populations cut off from one biotic zones. They state that a "belief that the another by water barriers). Darlington (1957) and present biota of the Southwest attained, and has Muller (1973) have accurately observed that it is retained, its present geographic distribution pq.or methodology to assume that the center of since the end of the Pliocene was based on cer­ range of a group or its center of greatest diver­ tain biogeographic inferences now largely in­ sity is necessarily its center of origin; therefore, validated by the Pleistocene fossil record. To be the use of number clues and center of range are, more specific, the fossil pollen record of at best, tenuous avenues of approach for deter­ Wisconsin-age pine parkland with spruce south mining a center of origin. This is the rationale for of the Colorado Plateau means sufficient down­ employing relative proportions of primitive and ward displacement of plant communities to derived species in any given area in conjunction allow several invasions of 'Arcto-Tertiary geo­ with geologic history, as well as the number floras' into the desert mountains during the clues. Pleistocene." Thus, I believe that the more highly From the Mexican-Central American center of derived S. aloeus, which is often sympatrically origin Strategus then extended its range north­ isolated with S. cessus on the more equable ward into the United States, southward into mountain islands in a sea of desert in the South­ South America, eastward into the Greater Antil­ west, also expanded northward from a Mexican les, and northward from South America into the refugium during pluvial periods of the Pleis­ Lesser Antilles. tocene when the Southwestern grasslands were invaded by pine-parkland. Isolation of the S. NORTH AMERICA aloeus line then occurred with the rapid recovery Howden (1966, 1969) has stated that most of post-glacial vegetation about 12,000 B.P. In North American scarabaeid genera were estab­ the American Southwest, then, there have been lished by Miocene times. Strategus cessatus at least two invasions of Strategus: once in the Wickham, the fossil from the Oligocene shale Miocene-Oligocene with the old S. cessus­ deposits at Florissant, Colorado, tends to sup­ cessatus line and again during the Pleistocene port this view. S. cessatus is very closely related with the S. aloeus line. The more optimum condi­ to S. cessus which occurs in the mountain chain tions of Louisiana, eastern Texas, etc., have from Durango in Mexico northwards into south­ permitted S. aloeus to become quite common ern Arizona and New Mexico. Strategus cessatus there. indicates that this group, expanding from the The two species in the southeastern United south, followed the Rockies as far north as the States, S. antaeus and S. splendens, have Antil­ 39th parallel during warmer Oligocene-Miocene lean affinities and possibly came from this area times. With the subsequent formation of the originally. There is good geologic evidence that major deserts in southwestern North America by a rather large land mass has been in existence in the mid-Pliocene (Axelrod, 1948, 1950; Cohn, northern Florida since the Middle Mi9cene 1965; Darrow, 1961), Strategus probably became (Clench and Turner, 1956; Howden, 1963; Hub- A REVISION OF THE GENUS Strategus / 159

bell, 1954; Vernon, 1951) that was not sub­ forest areas re-expanded during periods of merged during the Pleistocene (Altschuler and humid climatic conditions permitting the refuge Young, 1960). Goin (1958), Neill (1957), and Ross area populations to extend their ranges. Haffer (1965) have shown that there has been continu­ proposed that this rupturing and rejoining of the ous evolution influenced by recurrent isolation various forests was repeated several times and in this region from the Miocene to Recent time. led to a rapid differentiation of the Amazon forest The ancestors of these two species could have fauna in geologically very recent times. VUil­ arrived at this Floridian land mass at any time leumier (1971) has elaborated upon Haffer's since the Middle Miocene, but Young (1954) conclusions and shown that the chances were suggested Florida received the ancestors of the indeed very good for speciation in successive species of Antillean-Caribbean affinities during periods of ecological isolation: "These biologi­ the last part of the Pleistocene or during the cal data, combined with supportive geological W/irm interglacial periods, and this seems more evidence, show that climatic events during the ,:plausible in view of Antillean paleogeography. last million or so years have affected the biota of , During the Pleistocene glaciations Florida was South America as much as the Pleistocene gla­ undoubtedly serving as a refugium (Howden, cial changes affected the biotas of Eurasia and .1969), and S. antaeus and S. splendens, or their North America. Since most of South America lies immediate ancestors, expanded northward and within tropical latitudes, it is suggested here that ~estward from this area when the climate part of the diversity of species in the tropical ameliorated. I do not believe these ancestors areas of this continent is due to two historical originated in Mexico and moved east to Florida factors: the lack of wholesale elimination of during the interglacials as has been shown for species (compared with northern and high some prairie or semi-desert orthopterans (Hub­ latitudes), and ample opportunity for speciation bell, 1960). in successive periods of ecological isolation. The The presence of S. mormon in the southcen­ apparent paradox of the wealth of species in the tral United States might best be explained by a 'stable tropics' is partially explained by the fact westward movement of ancestral stock after in­ that the tropics have probably been quite unsta­ troduction into the southeastern United States ble, from the point of view of their biotas, during from the Caribbean. The development of mesic the Pleistocene and perhaps part of the Ter­ conditions could have then isolated the present tiary." These events undoubtedly influenced the surviving populations of S. mormon from its par­ evolution of Strategus in South America al­ ent stock. though to what extent is largely speculative due to the lack of certain aforementioned informa­ SOUTH AMERICA tion. Strategus probably spread into South America During the warmer interglacial periods of the east of the Andes during Tertiary times after the Pleistocene, the Amazon basin apparently be­ formation of the Panamanian land bridge. Post­ came a rather large inland sea when water levels ulating inferences for the historical dispersal of were raised about 50 m higher than today's level the South American members of the genus is (Haffer, 1969; Vuilleumier, 1971). I believe that highly problematical in view of the lack of addi­ this interglacial sea transgression into the Ama­ tional extensive specimen locality data, absence zon basin was directly responsible for the sub­ of fossils, and a lack of well-documented speciation of S. surinamensis into S. surinamen­ geologic history for some parts of the continent. sis hirtus south of the Amazon and S. Haffer (1969) has convincingly demonstrated surinamensis surinamensis north of the Amazon. that during the several dry climatic periods of the The interglacial inland sea barrier may also have Pleistocene and post-Pleistocene the Amazonian been an isolating mechanism for the ancestors forest was restricted to a number of smaller, dis­ of other South American species of Strategus, junct forests which served as refugia for numer­ and this might explain, in part, the seeming isola­ ous populations of forest animals which then dif­ tion of the species found exclusively south of the ferentiated from one another during these Amazon. The relative lack of Strategus species periods of geographic isolation. The isolated diversity in the Amazon region itself is indicative 160 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM of a re-colonization pattern from both the north today and that the more desert-like vegetation and the south. As Ross (1965) has noted, when which presently characterizes the low islands species or species groups of reasonably low va­ and the rain shadow areas of high islands was at gility have disjunct ranges, there is a good likeli­ times the dominant or even the only cover on hood that the disjunctions are the result of dis­ large islands: "Only high islands produce their ruptions by ecological changes in a once con­ own rain by adiabatic cooling, so only 01'\ high tinuous range. islands could conditions have been favorable for the persistence and continuing evolution of THE ANTILLES forest lizards during the periodic Pleistocene Accord'ing to Chace and Hobbs (1969) and droughts." I believe these conclusions apply Woodring (1954), conclusions on pre-Tertiary equally well to Strategus, and this might help to Caribb~an history are only tentative, but a explain their absence or presence on many of numberpf insular land masses probably existed the islands. in thei.Caribbean during the Cretaceous and, ex­ Simpson (1956) has succinctly noted that the cept' f9r a chain of volcanic islands in the area Lesser Antilles are a highly attenuated extension now encompassed by Cuba, most, if not all, of of the recent fauna of Trinidad and Venezuela; the extant islands, northern South America, and the 3-4 endemic species of Strategus repre­ parts~b_fCentral America were submerged during sented in the Lesser Antilles seem to bear this thispEir'iod. They state further that most of the out. Trinidad is a recently separated continental ::i~lands are probably not older than late island and is not considered to be part of the "Oligocene or early Miocene, and that there is no Lesser Antilles. S. a/oeus and S. surinamensis geological evidence to support the continuous surinamensis occur commonly in Trinidad and existence of land masses in the Caribbean prior on the mainland ten miles distant. The Lesser to the Eocene. Geologically and faunistically, the Antilles have probably never been connected Greater and Lesser Antilles have separate origins north of St. Vincent (Darlington, 1938) although and so will be discussed separately. all those islands within a single bank south of St. Vincent that are separated by shallow channels Lesser Antilles (Le., the Grenadines) were presumably joined The main Lesser Antillean chain has been during the last glaciation (15,000 B.P.) when sea emergent for no more than 11 million years (K-Ar levels were about 100 m lower than today (Yang dating), and Barbados probably emerged only V2 et al., 1974). Yang et al. have demonstrated that million years ago (Yang et al., 1974). Barbados is Anolis lizards are virtually the same within the not more than 300 m in elevation, receives less same bank of islands and different between rainfall than many of the higher islands, and is banks. Matthews (1966) has shown, with the not known to support any Strategus. Little is Scarabaeinae, that an immigrant pattern of dis­ known of Pliocene paleoclimates of the Carib­ tribution characterizes those species which are bean, but climates were known to be considera­ common to more than one island or to the conti­ bly drier during the low temperature periods of nent and one or more islands. Distance from the the Pleistocene glaciations. Bonatti and Gartner continent and island size are among the most (1973) have shown that these dry periods are important factors determining the number of compatible with other data indicating ice age species present on an island and, as a conse­ aridity in the western equatorial Atlantic as well quence, there is usually a gradual reduction in as in tropical South America. They suggested the number of species as one progresses out­ that one factor contributing to this phenomenon ward from the mainland. Matthews de­ was the decreased temperature of surface water monstrated that the Scarabaeinae of the Lesser in tropical and temperate oceans which would Antilles were clearly derived from South Ameri­ have significantly reduced the amount of water can forms or identical with them and that a fair vapor in the atmosphere above these areas. amount of divergence has occurred among the Using these data, Yang et al. (1974) convincingly outermost species. The species of Strategus in proposed that the rainforests of the high islands the Lesser Antilles display an identical pattern. were probably much less exten$ive then they are Grenada would be the most likely landfall for A REVISION OF THE GENUS Strategus / 161

rafting dispersal from the mainland; S. tar­ 1945a; Osborn, 1932; Pilsbry, 1930; Rivas, 1958; quinius is found here, and it is nearly identical Scharff, 1922; Schuchert, 1935; Schmidt, 1928), with S jugurtha and very similar to S. surinamen­ and (2) most of the fauna arrived by waif disper­ sis surinamensis in northern South America. sal from Central and South America (Baskin and Strategus verrilli is found on Dominica, and, Williams, 1966; Darlington, 1938, 1957; King, from all accounts, it is also very similar to the 1962; Koopman, 1958; Matthew, 1915, 1916, preceding three species. S. syphax occurs on 1918; Myers, 1938; Rosen and Bailey, 1963; Rui­ Guadeloupe and represents the northernmost bal, 1967; Simpson, 1956). The equally impress­ penetration of the Lesser Antilles from South ive arguments of the proponents of both factions America. S. syphax possesses several derived have been well documented and will not be re­ characters but, at the same time, retains some peated here. important ancestral ones which tend to indicate Although Heatwole and Levins (1972) and King perip,-heral divergence as noted by Matthews as (1962) provided valuable and significant data to wen as longer isolation from the parent stock. the idea of flotsam transport, I remain uncon­ The data seem to indicate secondary coloniza­ vinced that waif dispersal alone can account for tion at least up to Dominica because of the close the distribution pattern of Strategus in the phenetic (and presumably genetic) similarities Greater Antilles (Simpson, 1956, notwithstand­ between S. verrilli, S. tarquinius, S. surinamensis ing), but, on the other hand, there does not ap­ surf'namensis, and S. jugurtha as opposed to the pear to be any positive geologic evidence for a more primitive S. syphax. continuous land bridge. Therefore, rather than Strategus species have not been recorded advocate the conclusions of one camp over the from the three remaining large islands, Mar­ other, I have tried to utilize the reasoning of both tinique, St. Lucia, and St. Vincent. This is not approaches so as to best explain the current dis­ surprising since there has probably been little tribution of Strategus in the Greater Antilles. collecting there, but I would be willing to guess Khudoleyand Meyerhoff (1971), in a rather de­ that these islands do contain Strategus, and, finitive work on the geologic history of the considering the high degree of insular specia­ Greater Antilles, have demonstrated that the vol­ tion in the Antilles, there is a good likelihood that canic history of these islands suggests in-place these species may be new. Ballou (1914) re­ development, that is, a period of long stability as ported that Strategus species do not occur on opposed to the younger, independent Lesser Antigua or St. Kitts. Antilles tectonic unit. They state further that the S. talpa has expanded into some of the Lee­ Miocene was a time of active vertical uplift and ward Islands from the east and is not part of the considerable basinal or coastal subsidence, and "normal" colonization sequence of the Lesser that this was the last time during which impor­ Antilles as previously discussed. More will be tant thicknesses of sediments accumulated on said regarding S. talpa later. the existing Greater Antilles islands, with the ex­ ceptions of west-central Cuba, possibly the Greater Antilles younger Cauto basin of southeastern Cuba, and The paleogeography of the Greater Antilles locally within the grabens of Hispaniola. After remains the subject of considerable controversy Miocene times most deposition was confined to among geologists and particularly biogeog­ present coastal and offshore areas-reefs, es­ raphers. Much of the fauna and flora of the tuaries, and river mouths. The distribution of Greater Antilles has unquestionable Central Miocene marine deposits shows that numerous American affinities. Attempts to explain the large and small islands were present on the site methods of animal dispersal to the Greater Antil­ of the modern Greater Antilles. However, none of les fall into two basic schools of thought: (1) the islands was as large as today's major islands most of the fauna came from Central America by until the middle or late Miocene when most of way of a land bridge along the Nicaraguan swell the existing land areas emerged. Khudoley and to Jamaica and Hispaniola (Allen, 1911; Anthony, Meyerhoff observed that the faunal and floral 1925, 1926; Barbour, 1914, 1916; Bond, 1933; similarities between Central America and the Dunn, 1932, 1934; Forbes, 1930; Martorell, Greater Antilles suggests that direct land con- 162 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

nections existed perhaps via western Cuba or via ' isolation from the source of continued gene the Cayman ridge and Nicaraguan rise. Most im­ flow; Strategus ob/ongus and S. anachoreta are portantly, they state that this connection was the only two species that do not fit the hypotheti­ possibly in the form of fairly closely spaced is­ cal pattern well. land "stepping stones." I do not believe that the Ruibal (1967) observed that if inter-island col­ concept of closely spaced island stepping stones onizations were an important factor in the evolu­ falls within the realm of the land bridge theory tion of a group, it is logical to expect more com­ PJ~! se, but it does reasonably enable the mon species between islands. The high level of mechanisms of a land bridge and waif dispersal endemicity and differentiation shown by to function. The major difficulty of time and dis­ Strategus in the Greater Antilles indicates a rea­ ta~ce with respect to delicate, dessication­ sonably long separation from the mainland as prane, or salt-sensitive animals rafting across well as among the islands and among them­ f Ji:irge water barriers is eliminated as sequential selves. Only S. ob/ongus occurs on three islands ." ;qolonization is now possible across smaller (Grand Cayman, Hispaniola, and Puerto Rico) bodies of water by rafting or being carried by while all the other species are strictly endemic to ,storm winds (see Darlington, 1938, for an in­ a single major island. 'depth discussion of the latter). The occurrence of S. at/anticus on San Sal­ "'Utilizing the island "stepping stone" theory to vador in the southern Bahamas is almost cer­ which I adhere means that Jamaica would be the tainly the result of its ancestors being fortuit­ first island invaded from the Central American ously rafted or windblown from Cuba, His­ mainland followed by Hispaniola and then Cuba paniola, or Puerto Rica where it has close rela­ and Puerto Rico (the linear sequence of disper­ tives in S. aenobarbus, S. ajax, and S. talpa. In sal of Simpson, 1956). According to Matthews spite of favorable currents for waif dispersal (1966) and Simpson (1956) the last two islands from the Greater Antilles, the Bahamas in gen­ would be the last to be affected by new invasions eral are probably not inhabited by Strategus be­ and so would preserve early elements for the cause they are too low and dry. longest time. This appears nearly to be the case The extreme similarity between S. aenobarbus with Strategus (see Tables 5 and 6 and Fig. 175). and S. ajax suggests a fairly recent separation Among the Greater Antillean Strategus, the and isolation of these two species. Both species Cuban S. sarpedon (1) is the most primitive, fol­ are most closely related to S. caymani and S. lowed in order by S. inermis (2) (Hispaniola), S. simson. talpa (2) (Puerto Rico), S. at/anticus (3) (San Sal­ S. ob/ongus of Hispaniola and Puerto Rico is vador), S. ajax (3) and S. symphenax (3) (Cuba), probably a recent introduction to Grand Cayman S. aenobarbus (3) (Hispaniola), S. simson (4) Island, arriving there by rafting or being blown by (Jamaica), S. caymani (5) (Cayman Islands), S. storm winds; currents and strom tracks for this anachoreta (5) (Cuba), and finally S. ob/ongus (6) region agree well with this hypothesis. The af­ (Puerto Rica and Hispaniola). The solid arrows in finities of S. ob/ongus to the Central American S. the figure indicate the probable island "stepping a/oeus and S. /ongichomperus present a some­ stones" route which is partially supported by the what confusing situation zoogeographically. geology of the region. Furthermore, Darlington S. talpa occurs in Puerto Rico, the Virgin Is­ (1938) has shown that faunal relationships lands, and on some of the Leeward Islands. The among the islands are definitely from Jamaica to Virgin Islands are geologically a part of the Hispaniola to Cuba and not directly from Greater Antilles and may have once been loosely Jamaica to Cuba, and that the Greater Antilles joined with Puerto Rico, and so it is not surpris­ are faunistically more closely related to one ing that S. talpa occurs there. On the other hand, another than is anyone of the islands to the the occurrence of S. talpa in the Leewards indi­ mainland. Figure 175 shows the proposed major cates either a flotsam-jetsam dispersal to these routes of dispersal, and that, in general, the most islands or else introduction by the activities of primitive species are indeed peripherally located man prior to 1790 (since the types were taken among the Greater Antilles due to their longer from St. Barthelemy and described in 1792). S. A REVISION OF THE GENUS Strategus / 163

talpa is closely related to S. aenobarbus (Cuba) tem espoused by numerical taxonomy and to the and S. ajax (Hispaniola), and its ancestors un­ classical system of intuitively derived recon­ doubtedly came from Hispaniola. structions. I believe the objectivity of this ap­ Strategus simson (Jamaica) and S. caymani proach and espeCially its repeatability enable the (Cayman Islands) are closely related and are results to be clearly substantiated and arrived at both highly derived, with S. simson occupying by other systematists. the "port of entry" island of the Greater Antilles ASSUMPTIONS dispersal route from the mainland. I suspect the ancestors of S. caymani originally came to the Strategus is considered to be monophyletic c'ayman Islands fairly recently from Jamaica by and, as such, includes all the species assumed to waif or windblown dispersal. A previous island have descended from the hypothetical ancestor. cOfilnection from Jamaica to Grand Cayman is As Cracraft (1973) notes, this type of approach nol-- possible because of the intervening, very only assumes a relationship of hypothetical di3ep Barlett trough. common ancestry but cannot recognize and . " ;, Bel kin (1962) believes that the Caribbean was identify specific ancestors. All of the conclusions an important center of evolution of new mos­ presented here, therefore, are based on neon­ quitoes and other organisms rather than a bar­ tological data, and these data consist of the der to dispersal and that during island formation morphological character states and the inferred ttl/rough fragmentation of a land area, great en­ evolutionary directions of those characters. vironmental stress would come to bear on the Primitive characters are termed plesiomorph­ reduced and isolated surviving populations. ous, and derived characters are apomorphous Under these possibly rigorous conditions new (after Hennig, 1966). "Evolutionary (primitive­ adaptive types would have an ideal opportunity derived) sequences of the character states are to become quickly fixed. Bram (1967) reported hypothesized and taxa are clustered on the basis that 47 of 61 American species of the subgenus of shared derived character states. There is gen­ Culex (Culex) (Diptera: Culicidae) are found in eral agreement among systematists of very dif­ the Caribbean which was apparently their main ferent persuasions that reliable evol utionary center of origin. The relict pattern of dispersal sequences can be constructed for many charac­ displayed by Strategus in the Caribbean is very ter states" (Cracraft, 1973). similar to that situation found in the Table 5 shows the presumed states for the 19 Scarabaeinae by Matthews (1966) who believed characters used in this analysis. The following that the high degree of endemicity of the combinations of parameters were used to ascer­ Scarabaeinae in the Antilles is a reflection, not of tain plesiomorphy and apomorphy: (1) Simple to special insular evolution, but of the continued complex-as a general trend (that is not without survival of an ancient fauna for which the islands exceptions), less d~rived speCies generally pos­ have become a refuge. sess simpler or less elaborate modifications in PHYLOGENY their morphological character states. Thus, the absence of horns is usually a less derived state than the presence of horns, and a Simple horn is INTRODUCTION generally less derived than a horn with a forked The following is a computer-assisted cladistic or modified apex; similarly, an excised clypeal reconstruction of the presumed phylogeny or apex is more derived than an entire (Simple) evolutionary history of the lineage of the genus apex, and an enlarged mandibular lobe is more b~sed on the tenets of Hennig (1966) and Brun­ derived than undeveloped lobes. It must be em­ din (1966, 1972). Cladistic relationships refer to phasized that this parameter cannot be used in the branching sequence among the species in a and of itself as incorrect conclusions will result. ~hYlogenetic tree (cladogram) regardless of any For instance, elytral and pygidial punctation (a !Ime scal.e or of phenetic similarities among the more complex state) is here considered to be ~?nomlc units. It is a logical, repeatable system more primitive than a smooth or impunctate w Ich offers an alternative to the phenetic sys- elytra or pygidium based on the information pro- 164 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

vided by the remaining parameter. In general, eral on the elytra and pygidium in the majority of however, the other characters used in this study the less-derived genera observed; in other tend to fit the pattern of simple to complex as it is words, the punctation of the mesosternum correlated with primitive to derived. (2) In-group would follow the same trend as the other puncta­ and ex-group comparisons-similar character tion characters, i.e., loss of punctures being states which occur in closely related groups are apomorphic. However, in-group and ex-group probably ancestral; this is based on the infer­ comparisons indicate that this is not the case. ence that different lineages arising from a com­ The partially punctate mesosternum is here con­ meA ancestor diverge with regard to their sidered a less-derived condition than a com­ characters; in other words, a shared or common pletely punctate mesosternum because the large character is ancestral. The genus Strategus be­ majority of oryctine genera share this partially longs to the tribe Oryctini which, along with the punctate state, and, more importantly, because ve~/closely related Pentodontini, contains just the more highly derived Dynastini show in­ over a hundred genera, and of these roughly 35 creased or complete punctation of the mesos­ ,are monotypic. Comparisons of Strategus ternum. In this case, the wider ex-group com­ character states were made with the following 35 parisons provided more information than would pentodontine and oryctine genera: Pentodon, the Oryctini alone. A1i~$onotum, Heteronychus, Metanastes, Pyc­ Fossil sequences could not be used in this noschema, Dilobderus, Coe/osis, Ligyrus, study because they do not exist. The single Philoscaptus, Oxyligyrus, Bothynus, Aphonides, Oligocene elytron available exhibits gross punc­ Papuana, Anop/ognathus, Aphonus, Euetheo/a, tation, but the single fossil by itself cannot reli­ Oxygrylius, Orizabus, Pime/opus, Dipelicus, ably prove that this state is primitive. The puncta­ Phyllognathus, Calicnemis, Thronistes, Pericop­ tion of the elytra is considered primitive in this tus, Cyphonistes, Xy/oryctes, Heterogomphus, analysis based on in-group and ex-group com­ Enema, Oryctes, Podischnus, Scapanes, parisons and commonness, and the fossil evi­ Semanopterus, Megaceras, Licnostrategus, and dence tends to corroborate this conclusion. Trichogomphus. The shared character states possessed by METHODS many of these genera and Strategus suggest what is pleisomorphous and apomorphous with Table 6 shows, in a conventional data matrix, regard to all the characters being used, with the the scoring of the characters used in the analysis possible exceptions of the tubercles on the head of this genus where 0 is the primitive state, and 1, and the punctation on the mesosternum. The 2, etc. indicate an increasingly derived condition. presence of tubercles on the head (like the pre­ The %D column in the table suggests, in a very sence of horns) would normally be considered a general way, the relative degree of primitiveness more complex and derived state, but the occur­ or derivedness of each taxonomic unit, and, as rence of tubercles in such a wide array of oryc­ Edmonds (1972) and others have observed, the tine genera also suggests that this condition frequency of synapomorphy is a valid measure of might be primitive. It was decided to call the pre­ inferred phyletic (cladistic) relationship. The %D sence of tubercles on the head a derived condi­ is arrived at by dividing the number of derived tion which most of the genera in this tribe have characters by the total number (19) of characters attained; this conclusion was based on a wider being employed. ex-group comparison between the tribe Oryctini The method of arriving at a phylogeny is that of and the less-derived tribes Hexodontini and Cyc­ Camin and Sokal (1965), but because of the size locephalini and with the more advanced Dynas­ of the data matrix and the subsequent opera­ tini. Similarly, a completely punctate mesoster­ tions required of the data, the three sequential num might probably be considered CLADN programs written in FORTRAN IV by plesiomorphous as compared to a partially Bartcher (1966) for estimation of cladistic rela­ punctate (loss of punctures) mesosternum be­ tionships were used and adapted to an IBM 360 cause of the commonness of punctation in gen- computer. The data were processed at the Lin- A REVISION OF THE GENUS Strategus / 165

coin Computing Facility of the University of Ne­ branches by one or two branching points closer braska Computer Network. to the base, and tries to rearrange the various "The proposed method does not weight branches into clusters that are more parsimoni­ characters equally in the construction of the ous in terms of the number of evolutionary cladogram, since compatible characters are pre­ steps" (Bartcher, 1966). ferred over those that are incompatible. Charac­ RESULTS ters with few states tend to be more compatible than those with many. Since evolutionary steps Table 7 shows the compatibility matrix result­ a~e equally weighted, those with more states will ing from CLADN1. Characters and patterns are be more heavily weighted. However, the weight­ listed at the left and upper margins of the table, ing procedure agrees with the principles of the right and lower margins list compatibilities numerical taxonomy (Sokal and Sneath, 1963); it and the extra evolutionary steps required for isa,utomatic and a posteriori, based on the entire rows and columns respectively. Although there ,a:ilailable evidence rather than on a priori or are several row characters lacking com­ .character-by-character weighting as employed patibilities, none of the characters was consid­ in' conventional phylogenetic procedures" (Ca­ ered "bad," and so all were retained for the min and Sokal, 1965). In view of the above state­ CLADN2 analysis. m~nts, I have some doubts as to whether charac­ The CLADN2 program resulted in a basically ters seven and eight should be maintained as non-parsimonious procladogram upon which separate characters or joined to form one multi­ CLADN3 is to act. The summary table at the end state character, but they are here retained as two of this program indicated that 161 evolutionary characters. It is assumed that the cladogram with steps were required in the procladogram. A goal the minimum number of evolutionary steps of the CLADN3 program is to reduce this number (most parsimonious) is the best inference of the of evolutionary steps by as many as possible in correct cladistic relationships. order to arrive at the most parsimonious solu­ "The first program, CLADN1, computes a tion. compatibility matrix from the original data ma­ Fig. 176 shows the proposed phylogeny for the trix. The compatibility matrix reveals which genus Strategus as generated by the CLADN3 characters provide 'good' patterns that are rela­ program. The level numbers of successive furca­ tively close to the presumed correct cladogram. tions are given at the left-hand margin, and the It also points out 'bad' characters which appear four-letter abbreviations for each taxonomic unit to be miscoded or not to fit the assumptions of are given at the top of the figure. Evolutionary the method and should, therefore, be removed changes in character states are shown on each from consideration before proceeding with the branch of the cladogram. The left number is the analysis. character number, and the right number indi­ "The second program, CLADN2, applies the cates the number of evolutionary steps required monothetic clustering procedure to the data for that character in the indicated branch. Suc­ matrix to yield a relatively parsimonious clado­ cessive stages in clustering of CLADN3 (in this gram as a basis for further studies. The monothe­ case four were needed) has reduced the total tic procedure is applied to a data matrix from number of evolutionary steps required from the which the 'bad' characters have been deleted. original 161 to 116. "The third program, CLADN3, takes as input The results in Fig. 176 agree fairly well with my the procladogram produced by the monothetic preconceived, intuitively derived conclusions procedure and by performing a number of opera­ which were based largely on phenetic evidence. tions it improves its structure to yield a simpler The cladogram illustrates species clusters in a or more 'parsimonious' tree. The following oper­ fashion which tends to receive support from the ations are performed: (1) Program removes all zoogeographical analysis and vice versa. It empty internodes, (2) Program moves all com­ should probably be pointed out that the mon steps of branches with a common origin to zoogeography was done independently and be­ the base stem of these branches, (3) It moves fore the phylogenetic analysis so that the resul- 166 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM tant phylogeny would not unduly influence any has never been taken east of the Apalachicola zoogeographic considerations. The cladogram River in Florida. also tends to support the premise stated earlier Strategus anachoreta.-TRINIDAD (Ballou, 1914; that the frequency of synapomorphy is indeed a Smythe, 1920). valid measure of cladistic relationship. Strategus antaeus.-UNITED STATES: MICHIGAN ERRONEOUS RECORDS (1): no data. NEBRASKA (2): Lancaster: Malcolm. S. antaeus is not established in either of these The following records are erroneous or ex­ states. trerTlely unlikely. In certain instances they may Strategus argentinus.-BOLIVIA (Endrodi, refl~ct mislabeling, inadvertent conveyance by man 10 the place of capture, or are the result of 1973b). I am not convinced of the correct deter­ being substantially displaced by storm activity mination of these specimens. (in ~~ich case, if proven, the data would be valu­ Strategus centaurus.-VENEZUELA: MERIDA (1): a.b(~ for zoogeographic considerations). Merida. Strategus cessus.-UNITED STATES: CALIFOR­ Str.ategus ajax.-BRAZIL: CEARA (1): Fortaleza. NIA (1): San Diego: Cuyamaca. COSTA RICA: ALAJUELA (1): San Carlos. JAMAICA: No data (1). Strategus hipposiderus.-BRAZIL: BAHIA (1): Bahia. Strategus aloeus.-CUBA: No data (2). UNITED STATES. CALIFORNIA (3): No data. FLORIDA (28): Strategus mormon.-UNITED STATES: UTAH (Saylor, 1946). Alachua (27): Gainesville; Orange (1): No data. These specimens by F. W. Walker are mis­ Strategus sarpedon.-BRAZIL: No data (1). labeled; they provide another example and cor­ Strategus syphax.-CUBA, GRENADA, HIS­ roboration to a similar case discussed by Wood­ PANIOLA (Blackwelder, 1944). ruff (1973: 38). Woodruff has informed me (per­ sonal communication) that this species probably Strategus jugurtha.-VENEZUELA: ARAGUA: La occurs in the panhandle and far west but that it Providencia (Martorell, 1939). TABLE 5 PRIMITIVE AND DERIVED CHARACTER STATES IN Strategus. Conclusions based on in-group comparisons. comparisons with related genera. commonness. and simple versus complex states.

No. CHARACTER PRIMITIVE DERIVED

(1) apex on clypeus entire emarginate or excised (2) tubercles on head reduced or obsolete distinct (3) middle lobe of mandible small large (4) setae above eye absent present (5) eye size interocular width less than 3.0 interocular width greater than 3.0 transverse eye diameters transverse eye diameters (6) punctation on mesosternum anterior half setigerously punctate entirely and setigerously punctate (7) development of anterior horn not devloped; tuberculate developed (8) apex of anterior horn simple forked (9) development of posterior horns not developed developed (10) punctation of elytral disc large. ocellate punctures present large, ocellate punctures absent (11) apex of elytra without setae setae present (12) shape of parameres simple angulate or spined (13) punctation of pygidium densely and coarsely punctate sparsely punctate (14) setae on pygidium absent present (15) body length less than 35 mm greater than 35 mm (16) post-humeral punctation present absent (17) color dark. piceous light, castaneous (18) punctation on base of pronotum present absent (19) elytral luster dull shining A REVISION OF THE GENUS Strategus / 167

TABLE 6 SCORING OF CHARACTER STATES IN Strategus and % frequency of derived states (%D). 0 = primitive; 1 + = increasingly derived 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 %D

adolescens o 000 0 o 0 o 0 o 0 200 2 0.316

aenobarbus 000 0 o 2 o 0 o o 0 0 0 0.368

o 0 0 0 o 2 o 0 o o 0 0 0 0.368

aloeus 2 o 202 o 0 2 002 0.648

anachoreta o 2 o 2 0 2 0 0 0 o o o 2 0.526

antaeus o 3 00202 o 0 o 2 o 0 2 0.526

argentinus 000 o 000 2 002 0.526

atlanticus o o o 0 0 0 0 000 o 0 0.368

caymani 000 o 2 o o o o o 0.526

centaurus 020 02020 0 0 002 000 2 0.368

cessus o o o 0 0 0 0 0 0 0 0 000 2 0.263

o o 0 0 0 000 000 2 002 0.316

fallaciosus o o 0 0 0 0 000 000 2 0.368

fascinus 2 2 o 0 o o 0 2 0.737

hipposiderus 2 0 o o o 0 o 002 0.579 howdeni o o o o 000 2 002 0.526

inermis 000 o 0 0 0 000 o 0 o 2 0.316

jugurtha o 2 0 2 o o o 2 0.737

longichomperus 2 0 o 2 0 2 o 0 002 0.631 mandibularis o 2 o 2 0 2 000 002 0.579 mormon 003 o o o 002 o 2 0.579 oblongus o 2 o 2 0 2 o 0 o 002 0.579

sarpedon 000 o 0 0 0 0 0 0 0 0 0 0 0 o 2 0.158 simson 000 o 2 200 o 000 0.474 splendens o o 0 000 o 0 002 o 2 0.421 surinamensis o 202 o 0 o 002 0.631 symphenax 000 o 2 0 2 0 0 0 o 000 2 0.368

000 o 2 020 0 o 0 000 2 0.368

o 0 0 0 0 0 2 o 0 o o 0 0 0 0.316 targuinius o 202 o 0 o 2 2 0.737 validus o o 0 o 2 0 2 o 0 o 002 0.474 168 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

TABLE 7 CLADN1 COMPATIBILITY MATRIX OF THE 19 CHARACTERS 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19

1 0 1 2 1 1 1 1 1 2 1 0 1 1 1 1 2 1 0 1 2 19

2 1 0 3 1 1 1 2 1 2 1 1 1 1 1 1 2 1 1 1 0 23

3 2 3 0 2 2 3 4 1 4 2 3 0 2 2 5 4 3 1 0 2 43

.4 1 1 3 0 1 1 2 1 2 1 1 1 1 1 1 2 1 1 1 0 23

5 1 1 2 1 0 1 2 1 2 1 1 1 1 1 2 2 1 1 2 0 24

6 1 1 2 0 1 0 2 1 2 1 1 1 0 0 1 2 1 1 1 3 19

7 2 2 5 2 2 2 0 2 3 2 2 2 2 2 3 4 2 2 4 0 45

8 1 1 1 1 1 1 0 0 2 1 0 1 0 0 0 1 1 0 2 6 14

9 225 2 2 2 120 2 2 2 2 2 3 4 2 3 o 42

10 1 1 3 1 1 1 2 1 2 0 1 1 1 1 1 1 1 1 1 0 22

11 0 1 1 0 1 0 1 0 1 0 0 0 0 0 1 1 0 0 0 11 7

12 1 1 0 1 1 1 1 1 2 1 0 0 1 0 1 0 1 0 2 5 15

13 1 1 3 1 1 1 2 1 2 1 1 1 0 1 1 2 1 1 2 0 24

14 1 1 2 1 1 0 2 0 2 1 0 0 1 0 1 2 1 0 0 6 16

15 1 1 3 1 1 1 2 1 2 1 1 1 1 1 0 2 1 1 2 0 24

16 2 2 6 2 2 2 4 1 4 0 2 0 2 2 2 0 2 2 0 3 37

17 1 1 3 1 1 1 2 1 2 1 1 1 1 1 1 2 0 1 2 0 24

18 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 18 0

19 2 2 2 2 2 2 4 2 4 2 2 2 2 2 4 2 2 2 0 0 42

2 123 1 323 1 3 5 5 4 5 2 2 255

21 23 46 20 22 21 34 18 40 19 19 16 19 18 29 35 22 17 24 A REVISION OF THE GENUS Strategus I 169

ALPHABETICAL LIST OF VALID SPECIES AND SYNONYMS OF STRATEGUS

Valid Species Synonyms adolescens Kolbe, 1906. aenobarbus (Fabr.), 1775 = eurytus (Fabr.), 1775 = aenoburbns (Fabr.), 1787 (misprint) = eurytus (Fabr.), 1787 (redescription) = fossula (Beauvois), 1819 (new synonymy) = laterispinus Chapin, 1932b p.jax (Olivier), 1789 @/oeus (L.), 1758 = semiramis (Fabr.), 1801 (new synonymy) = aesalus (Laporte), 1840 = julian us Burmeister, 1847 (new synonymy) =piosomus Kolbe, 1906 (new synonymy) = roosevelti Casey, 1915 (new synonymy) = frontalis Casey, 1915 (new synonymy) = tarsalis Casey, 1915 (new synonymy) = gaiJ/ardi Casey, 1915 (new synonymy) anachoreta Burmeister, 1847 antaeus (Drury), 1773 = maimon (Fabr.), 1775 =maimon (Fabr.), 1787 (redescription) = divergens Casey, 1915 = atrolucens Casey, 1915 = pinorum Casey, 1915 = septentrionis Casey, 1915 = sinuatus Casey, 1915 = semistriatus Casey, 1915 = antaeus houstonensis Knaus, 1925 argentinus Kolbe, 1906 atlanticus, new species caymani, new speCies centaurus Kolbe, 1906 cessatus Wickham, 1914 (fossil) cessus LeConte, 1866 = beckeri Kolbe, 1906 (new synonymy) = cessus cavicauda (Casey), 1915 = durangoensis (Casey), 1915 = inflatus (Casey), 1915 = tantalus (Casey), 1915 craigi, new species fallaciosus Kolbe, 1906 fascinus Burmeister, 1847 hipposiderus, new species howdeni, new species inermis Arrow, 1947 jugurtha Burmeister, 1847 longichomperus, new species mandibularis Sternberg, 1910 mormon Burmeister, 1847 170 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

oblong us (Beauvois), 1807 = quadrifoveatus (Beauvois), 1807 = beauvoisi (new name; new synonymy) sarpedon (Burmeister,), 1847 simson (L.), 1758 = titanus (Fabr.), 1775 = simson (Fabr.), 1787 (redescription) = titanus (Fabr.), 1787 (redescription) splendens (Beauvios), 1809 = boscH (Beauvois), 1809 = cognatus (Casey), 1915 = carolin ensis (Casey), 1915 surinamensis surinamensis Burmeister, 1847 surinamensis hirtus Sternberg, 1910 = tridens Burmeister, 1847 (nomen nudum) I = kolbeanus Prell, 1934 (new synonymy) SY91phenax, new species ~yphax (Fabr.), 1775 = syphax (Fabr.) 1787 (redescription) = vulcan us (Fabr.), 1792 .ralpa (Fabr.), 1792 = barbigerus Chapin, 1932b (new synonymy) .tarquinius, new species Validus (Fabr.), 1775 = tricomis Jablonsky, 1785 .;, = validus (Fabr.), 1787 (redescription) = faunus (Bill berg), 1820 verrilli, new name

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Frey (eds.), The Quaternary of the 320. United States. Princeton Univ. Press, Princeton, New Jer­ Rosen, Donn Eric, and Reeve M. Bailey. 1963. The poeciliid sey, 922 pp. fishes (Cyprindontiformes), their structure, zoogeog­ Martorell, Luis F. 1939. Insects observed in the state of raphy, and systematics. Bull. American Mus. Nat. Hist. Aragua, Venezuela, South America. J. Agric. Univ. Puerto 126: 1-176. Rico 23(4): 177-232. Ross, Herbert H. 1965. Pleistocene events and insects. IN __. 1945a. A survey of the forest insects of Puerto Rico. Wright, Jr., H. E., and David G. Frey (eds.), The Quaternary Part 1. J. Agric. Univ. Puerto Rico 29(1): 70-354. of the United States. Princeton Univ. Press. Princeton. 922 __. 1945b. A survey of the forest insects of Puerto Rico. pp. Part 2. J. Agric. Univ. Puerto Rico 29(4): 355-608. Ruibal, R. 1967. Evolution and behavior in West Indian Matthew, William Diller. 1915. Climate and evolution. Ann. anoles. IN Milstead, William W. (ed.), Lizard ecology, a New York Acad. Sci. 24: 171-318. symposium. Univ. Missouri Press, Columbia. 300 pp. __. 1916. Supplemental note (See Barbour, 1916). Ann. Saylor Lawrence W. 1946. Synoptic revision of the United New York Acad. Sci. 27: 11-15. States scarab beetles of the subfamily Dynastinae, No.2: __. 1918. Affinities and origin of the Antillean mammals. tribe Oryctini (part). J. Washington Acad. Sci. 36(1): 16-21. Bull. Geol. Soc. America 29: 657-666. Schaeffer, Charles. 1915. New Coleoptera and miscellaneous 174 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

notes. J. New York Ent. Soc. 23(1): 47-55. Florida. Bull. Florida Geol. Surv. 33: 1-256. Scharff, Robert Francis. 1922. On the origin of the West In­ Verrill, A. Hyatt. 1906. Descriptions of two remarkable new dian Fauna. Bijdr. Tot de Dierkunde, Leiden 22:65-72. species of goliath beetles (Dynastes) from Dominica Is­ Schmidt, K. P. 1928. Amphibians and land reptiles of Porto land, Antilles. Brief contributions to zoology from the Rico, with a list of those reported from the Virgin Islands. museum of Yale University, no. LXVI. American J. Sci. (4th New York Acad. Sci., Sci. Surv. Porto Rico & Virgin lsI. series) 21(124): 317-320. 10(1): 1-160. __.1907. Description of a new species or sub-species of Schuchert, Charles. 1935. Historical geology of the hercules beetles from Dominica Island, B. W. I., with notes Antillean-Caribbean region. John Wiley & Sons, Inc., New on the habits and larvae of the common species and other York. 811 pp. beetles. Brief contributions to zoology from the museum Schl)feldt, R. W. 1884. Observations upon a collection of in­ of Yale University, no. LXVII. American J. Sci. (4th series) sects made in the vicinity of New Orleans, Louisiana, dur­ 24(142): 305-308. ing the years 1882 and 1883. Proc. U.S.N.M. 7(22): 331-338. Vuilleumier, Beryl Simpson. 1971. Pleistocene changes in the Simmonds, F. J. 1969. Biological control of sugar cane pests: fauna and flora of South America. Sci. 173: 771-780. a ,general survey. IN Williams, J. R., J. R. Melcalfe, R. W. Westwood, John O. 1837. Illustrations of exotic entomology, ;Mungomery, and R. Mathes (eds.), Pests of sugar cane. containing upwards of six hundred and fifty figures and EI~evier Publ. Co., Amsterdam. 568 pp. descriptions of foreign insects, interspersed with remarks Simpson, George Gaylord. 1956. Zoogeography of West In­ and reflections on their nature and properties. By Dru , .dian land mammals. American Mus. Novit. No. 1759: 1-28. Drury. A new edition, brought down to the present state of SJaane, Hans. 1725. A voyage to the islands Madera, Bar- the science, with the systematic characters of each bados, Nieves, S. Christophers, and Jamaica, with the species, synonyms, indexes, and other additional matter. : natural history of the herbs and trees, four-footed beasts, Vol. 1. Henry G. Bohn, London. 123 pp. fishes, birds, insects, reptiles, & c. of the last of those is­ Whitehead, D. C. 1964. Identification of p-hydroxbenzoid, ~ands; to which is prefix'd. an introduction, wherein is an vanillic, p-coumaric and ferulic acids in soils. Nature 202: account of the inhabitants, air, waters, diseases, trade & c. 417-418. of that place, with some relations concerning the Wickham, H. F. 1914. New Miocene Coleoptera from Floris­ neighboring continent, and islands of America. Illustrated sant. Bull. Mus. Compo Zoo I. 58(11): 423-494. with figures of the things described, which have not been Wolcott, George N. 1923. Insectae Portoricensis. A prelimi­ heretofore engraved; in large copper-plates as big as the nary annotated checklist of the insects of Porto Rico, with life. Vol. 2. British Museum, London. 205 pp. descriptions of some news (sic) species. J. Dept. Agric. Smythe, Eugene G. 1916. Report of the South Coast Labora­ Porto Rico 7(1): 1-312. tory. IN Fourth Rep. Board Commis. Agric. Porto Rico, __. 1926. Las plagas del cacao en Santo Domingo, y al­ Gov't. Porto Rico, Rio Piedras. 53 pp. gunas indicaciones para combatirlas. Rev. Agric. Puerto __. 1920. The white-grubs injuring sugar cane in Porto Rico 6: 11-12. Rico. J. Dept. Agric. Porto Rico 4(2): 3-29. __. 1936. "Insectae Borinquenses." A revised annotated Sokal, R. R., and P. H. A. Sneath. 1963. Principles of numeri­ check-list of the insects of Puerto Rico. J. Agric. Puerto cal taxonomy. W. H. Freeman and Co., San Francisco. 359 Rico 20(1): 1-600. pp. __. 1941. A supplement to "Insectae Borinquenses." J. Stahl, C. F., and L. C. Scaramuzza. 1929. Soil insects attack­ Agric. Univ. Puerto Rico 25(2): 33-158. ing sugar cane in Cuba. Tropical Plant Res. Found. Bull. __. 1948. Insects of Puerto Rico. J. Agric. Univ. Puerto No. 10: 1-19. Rico 32(1): 1-748. Staig, Robert A. 1931. The Fabrician types of insects in the __. 1950. The sugar-cane rhinoceros beetle. J. Econ. Ent. Hunterian collection at Glasgow University, part I. Col­ 43(3): 385. eoptera. Cambridge Univ. Press, Cambridge. 110 pp. Woodring, W. P. 1954. Caribbean land and sea through the Sternberg, Chr. 1910. Neue Dynastiden-Arten II. Ann. Soc. ages. Bull. Geol. Soc. America 65: 719-732. Ent. Belgique 54: 91-102. Woodruff, Robert Eugene. 1973. The scarab beetles of Torre-Bueno, J. R. de la. 1937. A glossary of entomology. Florida, part 1. Arthropods of Florida and Neighboring Brooklyn Ent. Soc., Brooklyn. 336 pp. Land Areas 8: 1-220. Valdes, Barry R. 1951. Control de gusanos blancos. Mem. Yang, Suh Yung, Michael Soule, and George Gorman. 1974. Asoc. Tecn. Azucareros Cuba 25: 51-56. Anolis lizards of the eastern Caribbean: a case study in Van Dinther, J. B. M. 1956. Insects of the coconut palm in evolution. 1. Genetic relationships, phylogeny, and coloni­ Suriname. Landbouwproefstation in Suriname Bull. No. zation sequence of the roquet group. Syst. Zool. 23: 387- 69: 1-27. 399. Vayssiere, P. 1965. Sur quelques insectes des palmiers en Young, Frank N. 1954. The water beetles of Florida. Univ. Amerique de Sud. Mededel Landouwhogesch Op­ Florida Studies, BioI. Sci. Ser. 5(1): 1-238. zoekingssta Ghent 30(3): 1571-1576. Zimsen, Ella. 1964. The type material of I. C. Fabricus. Vernon, R. O. 1951. Geology of Citrus and Levy counties, Munksgaard, Copenhagen. 656 pp. A REVISION OF THE GENUS Strategus / 175

I I I ,.. '" I ~ .... aenobarbus • ajax r:Y::~ns::~------I-- • atlanticus • caymani X simson

1 Fig. 1.-Distribution map for S. aenobarbus, S. ajax, S. at/anticus, S. caymani, S. simson.

~r------+------r--~ I ,I

-o d ""---.

• anachoreta X inermis • oblongu5 • sorpedon o symphenax 2 Fig. 2.-Distribution map for S. anachoreta, S. inermis, S. ob/ongus, S. sarpedon, S. symphenax. 176 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

,

syphax • talpa • tarquinius + verrilli o o

• ,J)

.----.....1 I) .• ',,, j

Fig. 3.-Distribution map for S. syphax, S. talpa, S. tarquinius, S. verilJi. A REVISION OF THE GENUS Strategus I 177

• antaeus .. spJendens

Fig. 4.-Distribution map for S. antaeus, S. sp/endens. Map copyright by University of Chicago, Dept. of Geography.

, / ~~----

I • aloeus (north of Canal Zone) --t-~------~--~~--t-~~~'-"->I

Fig. 5.-Distribution map for S. a/oeus (part). Map copyright by University of Chicago, Dept. of Geography. 178 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

0 cessatus (fossi I) cessus •A craigi howdeni • mormon •X fallaciosus I ------+----~-__l__---__:_I 6 Fig. B.-Distribution map for S. cessatus, S. cessus, S. craigi, S. fallaciosus, S. howdeni, S. mormon. Map copyright by University of Chicago, Dept. of Geography. A REVISION OF THE GENUS Strategus / 179

------

X hipposiderus o jugurtha • Jongichomperus

Fig. 7.-Distribution map for S. hipposiderus (part), S. jugurtha (part), S. /ongichomperus. Map copyright by University of Chicago, Dept. of Geography. 1S0 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

I

1 20 -- I

.. Fig. S.-Distribution map for S: a/oeus (part), S. argentinus, S. fascinus, S. validus. Map copyright by University of Chicago, Dept. of Geography. A REVISION OF THE GENUS Strategus I 181

centaurus •X hipposiderus 0 jugurtha ! I • mandibu la ris • 5. surinamensis .... 5. hirtus

Fig. g.-Distribution map for S. centaurus, S. hipposiderus (part), S. jugurtha (part), S. mandibuJaris, S. surinamensis sur· inamensis, S. surinamensis hirtus. Map copyright by University of Chicago, Dept. of Geography. 182 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

111111111111\11 II \ 11111111111111111111 ) 1 eM 1 2 10 eM 1 fa 11

IIIIInllllllljllili 14 1CM 1 2 15

17 Figs. 10--11.-8. adolescens (lectotype). Figs. 12-13.- S. aenobarbus (major development). Fig. 14.-8. aenobarbus (minor development). Figs. 15-16.-8. ajax (major development). Fig. 17.-8. ajax (minor development). A REVISION OF THE GENUS Strategus / 183

11111'111]1111 Plll[ 111111111111111111\ 1eM 1 II 18 1eM 1 :2 19

21

1'~~jllIll~lIJpll!1 22

Figs. 18-20.-S. aJoeus (North America). Figs. 21-22.-S. aJoeus (South America). 184 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

23

'111111111111\1111\ 26 / eM 1 2 27 Figs. 23-24.-8. aJoeus (minor development; South America). Figs. 25-26.-8. anachoreta (major development). Fig. 27.­ S. anachoreta (minor development). A REVISION OF THE GENUS Strategus / 185

llli pllllllllll1l1l ii Iii 1111\1111111111 j eM 1 2 28 IeM 1 2 29

111,/1111 111111111\ 30 1eM 1 2 31

J i i : 111111111111111"111 I I eM 1 2 32 I eM 33

34 Fig. 28.-5. anachoreta (minor development). Figs. 29-30.- S. antaeus (major development). Fig. 31.-5. antaeus (minor development). Figs. 32-33.-5. argentinus. Figs. 34--35.-5. at/anticus (holotype). 186 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

37

r

11111111111111 Ill) ) l' 'llljllll 11111'111\ eM 1 2 38 J eM 1 i 39

11111"1 II Ifjl 111\ 41 / CM 1 ~ Figs. 36-37.-5. caymani (major development; paratype). Figs. 38-39.-5. caymani (minor development; paratype). Figs. 40-41.-5. centaurus (major development). A REVISION OF THE GENUS Strategus / 187

11111111111111111111 1 eM 1 2 42

43

j11111111111111111111 44 eM 1 2 45

46 47 FigS. 42-43.-5. centaurus (minor development). Figs. 44-45.-5. cessus. Figs. 46-47.-5. craigi (holotype). 188 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

! III jllll\II11111\ 'I I eM 1 II 48

2 50 51

54 Figs. 48-49.-5. fallaciosus. Figs. 50-51.-5. fascinus (major development). Fig. 52.-5. fascinus (minor development). Figs. 53-54.-5. hipposiderus (holotype). A REVISION OF THE GENUS Strategus / 189

JlIl pllljllllP III/ IeM 1 2 55 56

11111'1 IIJlIjIlIlI lI/llftlllllll'llll 1 eM 1 2 59 IeM 1 2 60

nnp IBllmpllll J eM 11 I 61 Figs. 55-56.-5. howdeni (paratype). Figs. 57-58.-5. inermis (holotype). Figs. 59-60.-5. jugurtha (major development). Figs. 61-62.-5. jugurtha (minor development.) 190 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

64

ill 'I IIII1III '111111 lilli' ::,\;1111 1111\ j 1 eM 1 2 65 I eM 1 2 66 Figs. 63-64.-5. longichomperus (major development; para type). Figs. 6fr66.-5. longichomperus (minor development; paratype). A REVISION OF THE GENUS Strategus I 191

Iliil ill!III/l1l1J I CM 1 I 67

68

IlIIj 1IIIIlI! '1"111 "11 pili 1IIIIIIIll IeM, 1 l!: 69 / eM 1 ~ 70

Figs. 67-68.-8. mandibular/s (major development). Figs. 69-70.-8. mandibularis (minor development). Figs. 71-72.-8. mormon. 192 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

i eM 11 73

111'1111111111111111 11111'1111111[1111\ 1eM 1 2 77 1eM 1 II 78

I II' 1111111111111\ ! III jllll II \ III \ I \ IeM 1 2 79 IeM 1 2 80 Figs. 73-74.-5. obJongus (major development). Figs. 75-76. -5. obJongus (minor development). Figs. 77-78.-5. sarpedon. Figs. 79-80.-5. simson (major development). A REVISION OF THE GENUS Strategus / 193

llllj 1IIIjllli Jill I J / CM 1 It 81 82

11111" 11111111 JIll r ./ eM 1 2 83 84

IIIIPlllllllll'IIII 1eM 1 2 85 86

1111 II 111111111' 1111 1 CM 1 It 87 eM 88 Figs. 81-82.-S. simson (minor development. Figs. 83-84.-S. splendens. Figs. 85-86.-S. surinamensis surinamensis (major development). Figs. 87-88.-S. surinamenis surinamensis (minor development). 194 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

' II III' 11111111 II!TII 111111111 IIIIPlll\ j eM 1 !! 89 1eM 1 Ii! 90

1111'11111 IllIjllIll '1111111111111111"11 I eM 1 1I 1eM 1 Ii! 91 92

94

Figs. 89-90.-S. surinamensis hirtus (major development). Fi gs. 91-92.-S. surinamensis hirtus (minor development). Figs, 93-94.-S. symphenax (holotype). Figs. 95-96.-S. talpa (major development). A REVISION OF THE GENUS Strategus I 195

99

100

101 Fig. 97.-5. talpa (minor development). Figs. 98-99.-5. tarquinius (paratype). Figs. 100-101.-5. validus (major develop­ ment). 196 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

'lllllllllllIlljlllil 102 1CM 1 2 103

105

; 1111 jlllllll j I jlI11\ I CM 1 12 106 107 Figs. 102-103.-5. validus (minor development). Figs. 104-105.-5. syphax (major development). Figs. 106-107. (minor development). A REVISION OF THE GENUS Strategus / 197

... ' ... . .

109

111

113 115

119 116 117

1-- Figs. 108-123.-Lateral and caudal views of male genitalia. "'~109.-5. adoJescens. Figs. 110-111.-5. aenobarbus. Figs. 112-113.-5. ajax. Figs. 114-115.-5. aJoeus (North Amer.,,_ 116-117.-5. aJoeus (Colombia. Venezuela). Figs. 118-119.-5. anachoreta. Figs. 120-121.-5. antaeus. Figs. 122-123.- _ ·'~IIS. 198 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

125 127

129

131 133

Figs. 124-133.-Lateral and caudal views of male genitalia. Figs. 124-125.-5. at/anticus. Figs. 126-127.-5. caymanl. Figs. 128-129.-5. centaurus. Figs. 130-131.-5. cessus. Figs. 132-133.-5. craigi. A REVISION OF THE GENUS Strategus I 199

135

136 137

139

143 141

145 Figs. 134-147.-Lateral and caudal views of male gentalia. Figs. 134-135.-S. falJaciosus. Figs. 136-137.-S. fascinus. Figs. 136-139.-S. hipposiderus. Figs. 14D-141.-S. howdeni. Figs. 142-143.-S. inermis. Figs. 144-145.-S. jugurtha. Figs. 146- 147.-S.longichomperus. 200 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

151

149

153

155

157

159 161 Figs. 148-161.-Lateral and caudal views of male genitalia. Figs. 148-149.-S. mandibularis. Figs. 150-151.-S. mormon. 152-153.-S. oblong us. Figs. 154-155.-S. sarpedon. Figs. 156-157.-S. simson. Figs. 158-159.-S. splendens. Figs. 160- 161.-S. surinamensis surinamensis. A REVISION OF THE GENUS Strategus / 201

163 165

169 167

171 173

Figs. 162-173.-Lateral and caudal views of male genitalia. Figs. 162-163.-5. surinamensis hirtus. Figs. 164-165.-5. sym­ phenax. Figs. 166-167.-5. talpa. Figs. 168-169.-5. tarquinius. Figs. 170-171.-5. va/idus. Figs. 172-173.-5. syphax. 202 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

. .

174 Fig. 174.-Fossil elytron of Srategus cessatus Wickham (after Wickam 1914). A REVISION OF THE GENUS Strategus / 203

...

CD most primitive

@ most advanced

175 Fig. 175.-Proposed Greater Antillean dispersal routes of ancestral Strategus. Each number indicates the present occurrence of a species and its relative derived ness rating. 1 = most primitive; 5 = most advanced. Solid arrows = possible land connec­ tions or a closely spaced island "stepping stones" route. Dotted arrows = a probable flotsam dispersal route. 204 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

ATLA TALP.A£I'.O AJAX CAYM CENT SIMS SYF+1 SPLE f'.IORtv1 eRAI ADOL SARP INER CESS FALL SYMP ANte ANTA 1V\A.f\() HOM) ARGE HIPP VAll SURI FASC JJGU TARO OBLO LCN3 ALOE 4 16 181

11 11 41 81 ':1' 141 171

41 61 1 31 51 ' 131 141 31 16 1 L- 11 11 3 1 3 2 4 1 51 5 1 7 1 9 1 101 13 1 161 17 1 '---.-- L-

51 n 5 1 131 131 141 161 L...-r-- 2 1 21 3 2 3 1 4 1 61 7 1 7 1 7 1 81 91 9 1 101 101 111 131 131 141 151 161 191 '- L- '- '- 2 1 2 1 3 2 31 4 1 41 51 7 1 91 101 12 1 131 141 141 151 17 1 191 L- '---.--- '- 41 51 81 92 101 101 12 1 131 151 16 2 171 17 1 '---.--- '---r-- 11 2 1 4 1 51 51 6 1 7 2 7 2 81 91 151 151 7 1 191

131 T 191 I 176 Fig. 176.-Computer-generated cladistic reconstruction of the presumed phylogeny of the genus Strategus. See text for explanation. THe BOARD OF REGENTS Kermit Wagner, chairman O. Clinton Bellows Timothy R. Chappell James K. Say Kermit Hansen Ro~ert R. Koefoot, M.D. James H. Moylan Robert J. Prokop, M.D. Robert L. Raun Edward Schwartzkopf Robert G. Simmons, Jr. William F. Swanson, corporation secretary

THE PRESIDENT OF THE UNIVERSITY OF NEBRASKA

D. B. Varner

THE INTERIM CHANCELLOR OF THE UNIVERSITY OF NEBRASKA-LINCOLN

Adam C. Breckenridge

THE COMMITTEE ON SCHOLARLY PUBLICATIONS

Warren W. Caldwell, chairman David H. Gilbert, executive secretary Ned S. Hedges Royce Ronning Henry F. Holtzclaw PaulSchach Kenneth P. Pruess Gerald Thompson