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REVISION OF THE GENUS ISCHYRUS LACORDAIRE (1842) (EROTYLIDAE: TRIPLACINAE) OF NORTH AND CENTRAL AMERICA By PAUL EDWARD SKELLEY A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 1994 Copyright 1994 by Paul Edward Skelley ACKNOWLEDGMENTS I thank my committee, R. E. Woodruff and M. C. Thomas, Florida State Collection of Arthropods, and D. H. Habeck and J. W. Kimbrough, University of Florida, for their guidance, encouragement, assistance, and editorial comments for this and many other studies. Without them I would never have attained my goals. I thank M. A. Goodrich, Eastern Illinois University, for assistance during this study and for his continued support with other studies on the Erotylidae. For loans of specimens, including types, I thank the following people and their associated institutions: F. G. Andrews, California State Collection of Arthropods; R. S. Anderson, Canadian Museum of Nature; J. S. Ashe, R. W. Brooks, and R. A. B. Leschen, Snow Entomological Museum, University of Kansas; D. Azuma, Academy of Natural Sciences, Philadelphia; A. O. Bachmann, Museo Argentino de Ciencias Naturales; N. Berti, Museum National d'Histoire Naturelle, Paris; R. L. Blinn, University of Missouri; M. Brancucci, Naturhistorisches Museum Basel; R. L. Brown, Mississippi State University; C. Carlton, University of Arkansas; E. D. Cashatt, Illinois State Museum; R. D. Cave, Escuela Agricola Panamericana, Tegucigalpa, Honduras; D. S. Chandler, University of New Hampshire; F. A. Cholick, South Dakota State University; S. M. Clark, West Virginia Department of iii Agriculture; W. E. Clark, Auburn University; C. Covell, University of Louisville, Kentucky; D. P. Cowan, Western Michigan University; R. L. Davidson, Carnegie Museum of Natural History; K. Desender, Institute Royale des Sciences Naturalles de Belgique; M. A. Deyrup, Archbold Biological Station; M. E. Douglas, Arizona State University; O. V. Ferreira, FundaQ&o Oswaldo Cruz; R. W. Flowers, Florida A. & M. University; W. A. Foster, University of Cambridge, United Kingdom; P. H. Freytag, University of Kentucky; M. H. M. Galileo, Museu de Cidncias Naturais, Porto Alegre, Brazil; M. A. Goodrich, Eastern Illinois University; I. Goreyeb and T. P. Chaves, Museu Paraense Emilio Goeldi; L. H. Herman, American Museum of Natural History; M. W. Heyn, Clemson University; F. Hieke, Museum fOr Naturkunde der Humboldt- Universitdt zu Berlin; G. N. House, United States National Museum; N. Johnson and P. W. Kovarik, Ohio State University; D. H. Kavanaugh and R. Brett, California Academy of Sciences; M. D. Kerley, Natural History Museum, London; B. C. Kondratieff, Colorado State University; M. Kosztarab, Virginia Polytechnic Institute and State University; S. Krauth, University of Wisconsin; W. E. LaBerge and K. C. McGriffen, Illinois Natural History Survey; P. K. Lago, University of Mississippi; J. F. Lawrence, CSIRO, Canberra, Australia; R. Lawson, Chadron State College; R. E. Lewis, Iowa State University; J. K. Liebherr, K. E. M. Galley, and J. V. McHugh, Cornell University; J. McNamara, Canadian National Collection of Insects; J. E. McPherson, Southern iv Illinois University; O. Merkl, Hungarian Natural History Museum; C. R. Nelson, Brigham Young University; A. F. Newton, Jr., and P. Parillo, Field Museum of Natural History; D. NThez, Escuela Nacional de Ciencias Forestales, Honduras; M. F. O'Brien, University of Michigan Museum of Zoology; C. A. Olson, University of Arizona; G. Onore, Pontificia Universidad Cat6lica de Ecuador; C. S. Parron, North Carolina State University; S. Pratt, Museum of Comparative Zoology, Harvard University; A. Provonsha, Purdue University; B. C. Ratcliffe, University of Nebraska State Museum; E. G. Riley, Texas A. & M. University; R. A. Ronderos, Universidad Nacional de La Plata, Argentina; S. Santiago, Universidad Nacional Aut6noma de M6xico; G. Scherer, Zoologische Staatssammlung, Minchen; Y. Sedman, Western Illinois University; D. Shpeley and D. A. Pollock, Strickland Museum, University of Alberta; C. L. Smith, University of Georgia, Athens; R. R. Snelling, Los Angeles County Museum; C. A. Springer, Hastings College, Nebraska; F. W. Stehr, Jr., Michigan State University; A. L. TerAn, Fundaci6n Miguel Lillo, Tucuman, Argentina; R. E. Woodruff and M. C. Thomas, Florida State Collection of Arthropods; R. S. Zack, Washington State University; L. Zerche, Deutsches Entomologisches Institut, Eberswalde-Finow. I thank the following private collectors for allowing me to study their collections: A. Allen, R. J. Barney, J. L. Carr, J. M. Cicero, E. J. Ford, S. M. Fullerton, D. H. Habeck, M. A. Ivie, D. H. Kavanaugh, P. K. Lago, R. A. B. v Leschen, R. W. Lundgren, S. McCleve, J. V. McHugh, R. F. Morris, G. H. Nelson, T. K. Philips, R. Prange, W. Suter, R. H. Turnbow, Jr., J. E. Wappes, and J. Watts. For assistance in locating types, I thank W. A. Foster, Cambridge University, United Kingdom; F. Hieke, Museum fur Naturkund der Humboldt-Universitdt zu Berlin; D. S. Horning, University of Sydney; M. D. Kerley, Natural History Museum, London; G. C. McGavin, Oxford University. For assistance with the scanning electron microscope and specimen preparation techniques, I thank H. Cromroy and W. Carpenter, University of Florida. For darkroom assistance and other photographic reproductions, I thank J. Lotz, Division of Plant Industry, Florida Department of Agriculture and Consumer Services. I thank E. Mayr, J. McCarthy, and E. Langosy, Harvard University, for the support of an Ernst Mayr Grant, which allowed me to study specimens in both the Crotch Erotylidae Collection, Cambridge University, and the Natural History Museum, London. This study would not have been completed without their assistance. I thank M. C. Thomas, Florida State Collection of Arthropods, for assistance with computer graphic programs used in the illustrations. I thank all of the personnel (too numerous to mention) at the Florida Department of Agriculture and Consumer Services, Florida State Collection of Arthropods, for their support and tolerance during my eight years of study. If I have unknowingly omitted anyone, I thank them here for their assistance. Last, yet foremost, I thank my parents, Paul F. and Antoinette, and my wife, Lucy, for their encouragement and love over these many years. vii TABLE OF CONTENTS ACKNOWLEDGMENTS .................................. ........ iii ABSTRACT ................... ............................ x INTRODUCTION ............................................... 1 General Introduction ................................ 1 History ............................................. 2 Nomenclatural Status ............................... 6 Format of Species Accounts ......................... 8 Characters and Terminology ........................... 11 Basics .......................................... 11 Eye Facets ... Punctation ... Surface ...... Body Shape ... Color Pattern Head ......... Elytra ...... Ventral Lines Prosternum ... Mesosternum Metasternum First Visible Abdominal Sternite .............. Male Genitalia ................................ Female Genitalia .............................. Materials and Methods .............................. Specimens ..................................... Locating Types ................................ Collecting .................................... Equipment ..................................... Techniques .................................... Color Pattern Problems ........................ Rules of Thumb ................................ Results ....................................... ISCHYRUS LACORDAIRE ..................................... ARTIFICIAL KEY TO SPECIES ............................... SPECIES ACCOUNTS ........................................ Ischyrus aleator Boyle ............................. Ischyrus angularis Lacordaire ...................... Ischyrus auriculatus Lacordaire .................... viii ................................. ................................. ................................. ................................. ................................. ................................. ................................. ................................. ................................. ... .. .. ... .. .. ... .. .. .. Ischyrus bogotae Crotch ........................... Ischyrus boucardi Crotch .......................... Ischyrus chacojae Gorham .......................... Ischyrus collatinus Crotch ........................ Ischyrus distinguendus Lacordaire ................. Ischyrus dunedinensis Blatchley ................... Ischyrus elegantulus Lacordaire ................... Ischyrus ephippiatus Gorham ....................... Ischyrus episcaphulinus Gorham .................... Ischyrus frontalis Lacordaire ..................... Ischyrus fulmineus Delkeskamp ..................... Ischyrus incertus Lacordaire ...................... Ischyrus insolens Crotch .......................... Ischyrus pictus Gorham ............................ Ischyrus proximus Lacordaire ...................... Ischyrus quadripunctatus (Olivier) ................ Ischyrus quadripunctatus quadripunctatus (Olivier) Ischyrus quadripunctatus chiasticus Boyle, New Stat Ischyrus scriptus (Olivier) ....................... Ischyrus scutellaris Gorham ....................... Ischyrus septemsignatus Gorham .................... Ischyrus tetrasticus Gorham ....................... Ischyrus tripunctatus Crotch ...................... Ischyrus undulatus Gorham ......................... Ischyrus vespertilio Lacordaire ................... Ischyrus n. sp. 1 ................................. Ischyrus n. sp. 2 ................................. Ischyrus n. sp. 3 ................................. Ischyrus n. sp. 4 ................................. Ischyrus n. sp. 5 ................................. Ischyrus n. sp. 6 ................................. Ischyrus n. sp. 7 ................................. Ischyrus n. sp. 8 ................................. Ischyrus n. sp. 9 ................................. Ischyrus n. sp. 10 ................................ Ischyrus n. sp. 11 ................................ Ischyrus n. sp. 12 ................................ Ischyrus n. sp. 13 ................................ Ischyrus n. sp. 14 ................................ Ischyrus n. sp. 15 ................................ Ischyrus n. sp. 16 ................................ . 99 104 109 115 120 125 131 135 142 146 151 157 164 171 175 183 191 us 200 203 214 219 226 232 236 240 245 249 252 256 259 261 264 269 273 276 281 287 289 293 298 303 APPENDIX A: ISCHYRUS SPECIES NAMES .................... 308 APPENDIX B: COLLECTIONS STUDIED AND CODENS ............. 317 APPENDIX C: SPECIMEN LABEL DATA ........................ 320 LITERATURE CITED ....................................... 348 BIOGRAPHICAL SKETCH ...................................... 362 Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy REVISION OF THE GENUS ISCHYRUS LACORDAIRE (1842) (EROTYLIDAE: TRIPLACINAE) OF NORTH AND CENTRAL AMERICA By Paul Edward Skelley April 1994 Chairman: Robert E. Woodruff Major Department: Entomology and Nematology This revision treats all known species of the genus Ischyrus Lacordaire occurring from Panama north. A key to the species, descriptions and illustrations of terms used, habitus and genitalic illustrations for all species, distribution maps, detailed species descriptions, complete synonymies, label data from specimens studied, and comparisons with similar species are presented to aid in the identification of specimens. A list of all names used in combination with Ischyrus and their current status is presented to prevent future homonyms. A total of 26, valid previously described species is addressed, nine new synonymies made, and 16 new species described. INTRODUCTION General Introduction The family Erotylidae is composed of fungus-feeding beetles that vary in body size and color; many are elaborately patterned. Recently published catalogs detail the Old World species (Delkeskamp 1981; ChQj6 & ChQj6 1988, 1989, 1990). In contrast, Crotch's (1876) world revision of the Erotylidae was the last study to cover the entire New World fauna, but it provided no keys or illustrations and few descriptions. Since Crotch's time, the New World erotylid fauna has been studied intermittently, with scattered regional studies, catalogs, and species descriptions. Most New World erotylid genera have no modern revision. The purpose of this study is to begin bringing the nomenclature of the New World Erotylidae into the 20th century and to publish the results in a manner that will enable entomological students to identify their specimens. "Progress in Natural History necessarily starts from a basis of species, and until these are accurately described so that others can arrive at a knowledge of them no great advance is possible" (Horn 1887:7). Nomenclatural changes and lectotype designations made in this dissertation are not to be considered valid until 1 2 they are published. Holotypes are not selected and names are not proposed for the new species to avoid potential nomenclatural problems. History The first described member of the genus Ischyrus Lacordaire is Erotylus quadripunctatus Olivier (1792). The first use of the name Ischyrus was by Chevrolat in fascicle 5 of the second edition of Dejean's catalog (1836). Because most of the second edition was burned in a fire, the third edition of the Dejean catalog (1837) was immediately printed. (See Madge, 1988, for dates of publication and history of the Dejean catalogs.) Dejean's (1836, 1837) catalogs were among the first works to split the genus Erotylus Fabricius (1775) into more manageable, related taxa. Species and generic concepts were in their infancy during the early 1800s. For the Erotylidae, Dejean's genera appear to be groups of species with similar body size and color. This is evident in the proposed genus Ischyrus Chevrolat containing large dull-black species, Mycotretus Chevrolat containing smaller species with yellow and black color patterns, and Lybas Chevrolat containing species that were oval and solid red. Dejean's catalogs were simply checklists of his collection in which many generic and specific names were proposed. Because the Dejean names lacked descriptions, they were often ignored by early taxonomists. The next use of the name Ischyrus was by Falderman (1837) in describing Ischyrus lepidus, presently a member of Triplax Herbst (1793) (fide ChOj6 & ChOj6 1990). The name Ischyrus was then used as a subgroup of Erotylus Fabricius (1775) by Gu6rin-M6neville (1841) in describing E. (Ischyrus) nebulosus, presently a member of Brachysphaenus Lacordaire (1842) (fide Crotch 1876). These uses of "Ischyrus" illustrate early workers' attempts at placing new species into taxa based on superficial characters. Lacordaire (1842) was the first person to provide a description of the genus Ischyrus, for which he gave credit to Chevrolat in Dejean (1836) as the author of the name. He moved many previously described species into Ischyrus and described 17 new species. Unfortunately, Lacordaire did not designate a type species for Ischyrus. Lacordaire split the genus into two divisions. The first division contained species with larger body size and strongly keeled prosternum, mostly species listed in the Dejean catalogs as Ischyrus Chevrolat. The second division contained species with smaller body size and a weakly keeled prosternum, mostly species listed under Mycotretus Chevrolat in the Dejean catalogs, including Erotylus quadripunctatus Olivier. Lacordaire based his generic concepts on reasonably sound morphological features that are still in use. He also maintained many previously published species names that had not been described [nomina 4 nuda]. With his descriptions, Lacordaire became the author of these names. Subsequent authors, Guerin-Meneville (1844) and Erichson (1847), used Lacordaire's generic names and described seven additional species of Ischyrus. Crotch (1873b) separated Lacordaire's divisions into distinct genera. The first division, containing species of larger size, he named Megischyrus. The second division, containing species of smaller size, he maintained as Ischyrus, crediting Lacordaire as the describer and noting the original use of the name by Chevrolat in Dejean (1836). At that time, it was common to give credit for a name to the first describer of the taxon and not to the author of that name. Following this trend, Crotch cited it as Ischyrus Lacordaire. In 1873 Crotch received a grant to visit tropical Australia and adjacent islands to collect natural history specimens. Before his departure from Cambridge, U.K., he placed his collection and manuscript in the care of E. W. Janson. Unfortunately, Crotch became ill and died in 1874 while in the U.S.A. Edward W. Janson put Crotch's manuscript into final form and published it. Numerous notes within the revision are undoubtedly those of Janson. Without Crotch's input, many of the species problems alluded to within the text were left unresolved. This "revision" is simply an annotated catalog of the species previously described, with brief descriptions of the new species. Yet, it has been the basis for all subsequent studies of the family. The name Ischyrus Lacordaire, as cited and used in Crotch's revision, was accepted and used by all subsequent workers, with the exception of Alvarenga (1965, discussed below). Crotch (1876) listed 44 species of Ischyrus, 13 of which were new species. Ischyrus Lacordaire (sensu Crotch) has been referenced in a multitude of papers and texts. Many of these are simply species descriptions, regional studies, catalogs, general entomological texts, or biological accounts. These are too numerous to list here, but they can be found in the species accounts of this revision. Some of the more important references are worth noting. Gemminger and Harold's (1876) Catalogus Coleopterorum was published shortly after Crotch's revision and contains the names as used by Crotch. Gorham (1887- 1899), in the Biologia Centrali-Americana, gave accounts for 22 species of Ischyrus that included 10 new species. Kuhnt (1909, 1911) provided two catalogs, the first of which contained a simple grouping of the species by various color pattern characters. The most recent catalog of the genus Ischyrus Lacordaire is Blackwelder's (1945) Checklist of the Coleopterous Insects of Mexico, Central America, the West Indies, and South America, which lists 56 names. Boyle (1954, 1956) revised the family Erotylidae for America, 6 north of Mexico, thoroughly describing the genus Ischyrus and one new species. The most recent paper of importance to the genus Ischyrus Lacordaire is Alvarenga (1965). Following current nomenclatural rules, Alvarenga realized that Crotch (1873b) had incorrectly applied the name Ischyrus and selected an invalid type species. Alvarenga made some major changes in the standing of the names Ischyrus and Megischyrus, which are discussed in the following section. Before the present study, there were 65 valid species placed in the genus Ischyrus Lacordaire. This study addresses 26 of these, synonymizes 9, and describes 16 new species. This brings the total to 72 species. (See Appendix A for a list of all specific names used in association with Ischyrus and their current status.) Nomenclatural Status The early nomenclature of this genus is typical for many taxa, with names being proposed and ignored. The name Ischyrus Chevrolat in Dejean (1836) is a nomen nudum (name without description) and was not valid in the opinion of some early workers. Thus, the first use of the name to be accompanied by a description was the valid name. Crotch (1873b) undoubtedly had this in mind when he raised Lacordaire's divisions to full generic status. Crotch proposed the name Megischyrus for Lacordaire's first division, designating Erotylus undatus Olivier as the type species. Crotch retained the name Ischyrus for the second division, crediting Lacordaire as the first describer of the genus, and designated Erotylus quadripunctatus Olivier as the type species. This was accepted and used by all subsequent workers until Alvarenga (1965) found some errors in the type designations and credits for these genera. According to the current International Code of Zoological Nomenclature (1985) the first valid use of a name is to be followed--the Law of Priority. Before 1931, the first use of a generic name is valid if it is followed by a description or has a valid species name listed under it, an indication. Also, in designating a type species for a genus subsequent to its first use, the reviser designating the type must choose a species from those originally included within the genus (as indicated by Barber & Bidwell 1940). This makes the Dejean catalogs (1836, 1837) the first valid uses of the generic name Ischyrus, and the type species should have been chosen from the species listed within it. According to these rules, Crotch (1873b) had incorrectly chosen the type species of Ischyrus, because Erotylus quadripunctatus was not listed in Dejean (1836) as Ischyrus. Erotylus undatus Olivier was listed under the name Ischyrus Chevrolat in Dejean (1836). Crotch chose Erotylus undatus as the type species of his genus Megischyrus. This makes Megischyrus Crotch an objective synonym of Ischyrus Chevrolat, leaving Ischyrus Lacordaire (sensu Crotch) without a valid name. 8 Discovering these problems, Alvarenga (1965) corrected them by synonymizing Megischyrus Crotch under Ischyrus Chevrolat, designating Erotylus undatus Olivier as the type species. For Ischyrus Lacordaire (sensu Crotch) he proposed the name Micrischyrus, designating Erotylus quadripunctatus Olivier as the type species. In essence, he discarded one name, moved another, and proposed a third. In my opinion, Alvarenga's actions were not fully justifiable because they create unnecessary confusion. The main purpose of the International Code of Zoological Nomenclature is to stabilize nomenclature, as stated in its preamble. The code has provisions (Article 79) to conserve long-standing, widely accepted names like Megischyrus Crotch and Ischyrus Lacordaire (sensu Crotch). Therefore, a proposal has been submitted to the International Commission of Zoological Nomenclature (Skelley & Goodrich, in press) to conserve these names by suppressing all uses and indications of the name Ischyrus prior to Ischyrus Lacordaire (1842). Pending the ruling of the Commission, I am using the name Ischyrus Lacordaire (sensu Crotch), in preference to Micrischyrus Alvarenga, as the name for the genus studied in this revision. Format of Species Accounts Title. This heading is the valid name for the species to be discussed. Synonymy. This section lists all combinations of the specific name and any synonyms of the valid name. These are organized in the following manner: Genus species describer date:page [comments], author of the synonymy or combination date:page. Example: Engis variegata Dejean 1821:45 [nomen nudum], Gemminger & Harold 1876:3691. Diagnosis. This section lists the set of characters separating the species under consideration from all known species in the genus. Description. This section is a detailed description of the species being discussed. It is organized in the following manner: body measurements, overall appearance, color pattern, head and antenna, visible mouthparts, pronotum, scutellum, elytra, prosternum, mesosternum, metasternum, first visible abdominal sternite, male genitalia, female genitalia, stridulatory files, sexual dimorphisms. Legs are not described because no characters of use at the species level were found. Variation. This section is a discussion of the color pattern or morphological features that vary from specimen to specimen in the species. Type. Here I list the type specimens for all species names appearing in the synonymy, their label data, type locality (if not on their labels), current location, sex, and if they were studied. Label data for type specimens are given in the following manner: "/ label data/ [my comments] label data/" [XXXX]. The quotation marks, ", indicate the beginning and ending of the data. The slash marks, /, indicate the 10 beginning and ending of an individual label. My comments are placed in brackets, [ ], and usually indicate paper color or label shape. When no comment is made, the paper is white. After the label data are presented, a coden is given in brackets, [XXXX], for the institution (see Appendix B) where the specimen it is currently located. Many species needed lectotypes designated. A red "lectotype" label was placed on these specimens and dated when they were designated as the "type." Type specimens of previously described species were dissected only if it was absolutely necessary to solve a species problem. Since this was such a rare occurrence, determining the sex of the types was accomplished using external characters. In some cases, where sexual dimorphism is distinct, the determination was easy. In most cases, when the sex was unclear, it is listed as "not determined." Specimens examined. Here I list the label data for all specimens studied, if there were fewer than ten records. Species with more than ten collection records have their data presented in tabular form in Appendix C to help keep the text uncluttered and to present it in a more usable format. Distribution. A brief account of the known geographic distribution of the species is presented. A map is provided for each species to illustrate the distribution. Etymology. An attempt is made to present what the species epithet means and possibly why the original author chose it. This is included more for historical purposes and may help the reader remember the name. Brown's (1985) Composition of Scientific Words was an indispensable reference in this research. Taxonomic notes. This section includes nomenclatural aspects that need clarification. I present some unsolved problems of taxonomic importance indicating that this revision is just a beginning. Biologv. This section is included only for the few species where some biological or life history data are known. Remarks. This section contains statements on similar species and how to separate them from other species being covered. References. This section lists the references where the species name occurs. Each species name appearing in the synonymy is listed separately to aid in future literature searches for specific names other than the valid name. Characters and Terminology Basics Most general terms follow the meaning presented in The Torre-Bueno Glossary of Entomology (Nichols & Schuh 1989). Terms for the genitalia follow Sharp and Muir (1912), 12 Tanner (1927), and Boyle (1956). Some characters and terms need further explanation, as follows. Eye Facets Eye facet size, coarse vs. fine, is used to separate many genera in the Triplacinae. This character is based on the relative size and distinctiveness of the eye facets in relation to the head. Coarse facets (Fig. la) are larger and more prominent, often bulging from the surface. Fine facets (Fig. Ib) are smaller and less prominent, with a smoother eye surface. There are many species that are intermediate in facet development, making this character difficult to interpret. Punctation The puncture size is compared with the eye facet: facet diameter to puncture diameter. Most Ischyrus species have coarse punctation where the punctures are as large or larger in diameter than a facet. Punctures can be normal, impressed, or foveate. A normal puncture appears like a simple pinprick. Impressed punctures are deep punctures, usually with a rounded edge and bottom. Foveate punctures are large, shallow, flat- bottomed punctures, usually with a distinct edge. Surface The body is generally covered with a hexagonal micro- sculpturing. Variations in the strength of this microreticulation change the surface from shiny to dull and are visible to the naked eye. Figure 1. Dorsal view of heads showing eyes: a.) Ischyrus q. quadripunctatus (Olivier) with coarsely faceted eyes, line = 0.5 mm; b.) Tritoma atriventris LeConte with finely faceted eyes, line = 0.38 mm. The arrow points to the ocular stria. Body Shape Most of the species are flattened to slightly convex dorsally; vespertilio Lacordaire and duponti Lacordiare are convex dorsally (Fig. 2a, 2b). Most parallel-sided bodies (Fig. 2c) are flattened dorsally. Most elongate bodies have the sides parabolically rounded (Fig. 2d) and are slightly convex above. Oval bodies (Fig. 2e) are rounded on the sides and can be flattened or convex dorsally. Ovoid bodies are egg-shaped (Fig. 2f) and wider anteriorly and can be flattened or convex dorsally. Color Pattern Color patterns are the most useful characters in the recognition of species. Understanding the terms for various aspects of the color pattern is essential in using the key (Fig. 3). A band is a wide transverse marking, wider than long. A stripe is a longitudinal marking, longer than wide. A spot is a small marking, usually circular or elongate. I use the term "fascia" to mean a band that may be broken by additional markings. A free spot is not connected to any margin. A tooth-like spot is a triangular spot connected to a margin. The color patterns of most species are variations of a basic pattern. I have named the components of the pattern in reference to its position on the body (Fig. 4): anterior pronotal, free pronotal, basal pronotal, pronotal hind angle, humeral, subhumeral, scutellar, elytral suture, c d e f Figure 2. Ischyrus body shapes: a.) flattened; b.) convex; c.) parallel-sided; d.) elongate; e.) oval; f.) ovoid. st FSp B {B St Fj F Figure 3. Diagrammatic representation of color pattern terms: B = band, F = fascia, FSp = free spot, Sp = spot, St = stripe, TSp = tooth- like spot. Figure 4. Diagrammatic representation of a generalized color pattern indicating spot names: Ae = apical elytral, Ap = anterior pronotal, Bp = basal pronotal, Ce = central elytral, Es = elytral suture, Fp = free pronotal, Hu = humeral, Le = lateral elytral, Pha = pronotal hind angle, Shu = subhumeral, Sc = scutellar, Sc + Hu = basal elytral band, Ce + Le = central elytral band. 18 central elytral, lateral elytral, and apical elytral. When the scutellar and humeral spots are connected, they make up the basal elytral band. When the central and lateral elytral spots are connected, they make up the central elytral band. Head The dorsal surface of the head has relatively few useful characters. On each side of the vertex following the margin of the eye is a line or shallow groove, the ocular stria (Fig. 1). Its length appears to be useful at generic levels and is commented upon for future reference. Since this line is shorter than the eye length in the vast majority of Ischyrus, its length is given as a decimal that indicates how far forward the line reaches on the eye. For example, it could reach "0.75 distance to the anterior angle of the eye," which means the line stops 3/4 of the distance from the base to the anterior angle of the eye. In Figure la, the ocular stria stops at the anterior angle. The head size vs. eye size is given as a proportion: head width between eyes = "N" eye widths (Fig. 5). The smaller the eyes or the wider the head, the larger the number "N". The base of the head often has structures which have been called stridulatory files (Alexander et al. 1963; Arrow 1924, 1925, 1942; Delkeskamp 1959). These structures appear as iridescent spots under a dissecting microscope. Study with the scanning electron microscope shows them to Figure 5. Diagrammatic dorsal view of a head showing the measurements used for determining the ratio of head to eye width; dorsal head width between eyes = y/x times eye width. 20 be patches of parallel ridges. Even though these have been called stridulatory, I have found no statement about the sound produced. They appear only on males in some species and on both sexes in others. The exact function of these structures is unknown. Representative species' antennae are illustrated on a single plate for ease in comparison (Fig. 6). If a species antenna is not illustrated, a reference is given to the antenna that is most similar: "(similar to Fig. X)", where "X" is the figure number. The shape and proportions of the maxillary and labial palp terminal segments are important for species distinctions. These are illustrated on a single plate (Fig. 7) for ease in comparing shapes and proportions. This plate illustrates the palpi of representative species to show the basic forms. The descriptions of species with similar palpi have the statement "(similar to Fig. X)," which refers to the figure "X" that is most similar to that seen in the species being discussed. The "triangular mentum" is a characteristic of Ischyrus. The "triangle" is a sunken area of the mentum surrounded by a ridge (Fig. 8). The triangular sunken area is the mental plate. The ridge surrounding the plate is the mental ridge; it is often extended forward at the middle as a sharp divider between the labial palps. This projection is called the medial ridge extension. As with palp terminal segments, representative menta are b a g c 1 e f I I n m Figure 6. Antennae, line = 1.0 mm: a.) Ischyrus distinguendus Lacordaire; b.) I. n. sp. 2; c.) I. insolens Crotch; d.) I. bogotae Crotch; e.) I. angularis Lacordaire; f.) I. q. quadripunctatus (Olivier); g.) I. n. sp. 3; h.) I. n. sp. 4; i.) I. n. sp. 6; j.) I. n. sp. 5; k.) I. n. sp. 1; 1.) I. n. sp. 9; m.) I. n. sp. 14; n.) Megischyrus zonalis (Lacordaire); o.) Megischyrus sp. 7)? e 72 1 P f g o k n q >7 Figure 7. Terminal segments of labial palp (left) and maxillary palp (right), line = 0.33 mm: a.) Ischyrus distinguendus Lacordaire; b.) I. n. sp. 2; c.) I. insolens Crotch; d.) I. bogotae Crotch; e.) I. septemsignatus Gorham; f.) I. q. quadripunctatus (Olivier); g.) I. n. sp. 3; h.) I. undulatus Gorham; i.) I. n. sp. 4; j.) I. n. sp. 6; k.) I. n. sp. 7; 1.) I. n. sp. 5; m.) I. aleator Boyle; n.) I. n. sp. 1; o.) I. n. sp. 9; p.) I. ephippiatus Gorham; q.) I. tripunctatus Crotch. h 1 m 0 a b d e f Figure 8. Menta, line = 0.33 mm: a.) Ischyrus distinguendus Lacordaire; b.) I. n. sp. 2; c.) I. insolens Crotch; d.) I. bogotae Crotch; e.) I. n. sp. 3; f.) I. n. sp. 4; g.) I. n. sp. 6; h.) I. n. sp. 5; i.) I. aleator Boyle; j.) I. ephippiatus Gorham; Lp = labial palp, P = mental plate, R = mental ridge, Re = mental ridge medial extension. illustrated on a single plate (Fig. 8) and are often referred to in the text as "(similar to Fig. X)." Elytra Except for color patterns mentioned above, the elytra have few useful characters. One is the number of elytral striae and the strength of the strial punctures. Most species have seven complete striae; stria I next to the elytral suture, striae V and VI originating at the humerus. Stria VIII on most species is reduced to a short row of punctures visible at the basal quarter under the humerus and/or the apical quarter near the lateral edge of each elytron. Two species (I. n. sp. 6 & I. n. sp. 7) have stria VIII complete, one species (I. n. sp. 1) has parts of stria IX visible. The size of the strial interval punctures is variable from species to species. In the majority of species the punctures are small and obscured in the microsculpturing. In a few other species they are large and distinct, occasionally obscuring the strial punctures. Ventral Lines A "line" refers to a ridge or fine groove on a sclerite that is not a suture, but probably has some supportive function. These lines surround the coxae and often extend onto the sclerite (Fig. 9). The term "coxal line" is used in reference to the line on the median side of the coxa. The lines anterior or posterior to the coxae are referred to by the structure they are on or near; e.g., Figure 9. Ventral view of a.) Ischyrus proximus Lacordaire and b.) I. n. sp. 1, line = 1.0 mm: Al = abdominal coxal line, Ee = epipleural fold of elytron, Lap = prosternal line anterior to procoxa, Lc = coxal line continuous around coxa, Lnc = coxal line not continuous around coxa, Lpm = metasternal line posterior to mesocoxa, Mp = mental plate, Msl = mesocoxal line, Mtl = metacoxal line, Pe = pronotal epipleuron, P1 = procoxal line, Pps = prosternal- pronotal epipleuron suture, Ps = prosternum, Psp = prosternal plate. the prosternal line in front of the procoxa or the metasternal line behind the mesocoxa. In the majority of Ischyrus species the coxal lines are not connected to other lines and extend onto the sclerite (Fig. 9a). If the coxal line is connected to the anterior or posterior line, it is "continuous around the coxae," because there is no break in the line surrounding the coxae (Fig. 9), and the line does not extend onto the sclerite. Prosternum The prosternum is a T-shaped sclerite with the base between the procoxae and the cap anterior to the procoxae. The anteromedial section of the prosternum is often keel- like and elevated above the sides to the level between the procoxae (Fig. 10), straight in profile. This elevation makes the anterior margin project at the middle, appearing as a "pitcher-like lip" (according to Boyle 1956). When the prosternum is keeled and has this "pitcher-like lip," I refer to it as being "pinched," because it appears laterally pinched. The strength of this pinch, also the amount it projects, is variable throughout Ischyrus and can be absent (Fig. 10a, 10b), weak (Fig. 10c, 10d), or strong (Fig. 10e, 10f). In a few species (for example, I. n. sp. 5, I. n. sp. 6, and I. n. sp. 7) the prosternal keel appears anteriorly swollen just behind the margin (Fig. 10g, 10h). I call this "swollen above the pinch." ' .e ,, .. c, riii~ 1 Figure 10. Prosternal development: a.) ventral and b.) lateral view of Ischyrus aleator Boyle [Mexico, Sonora] prothorax lacking anterior pinch, line = 0.60 mm; c.) ventral and d.) lateral view of Ischyrus q. quadripunctatus (Olivier) [USA, Florida] prothorax with weakly developed prosternal keel and pinch, line = 0.60 mm; e.) ventral and f.) lateral view of Ischyrus scutellaris Gorham [Mexico, Yucatan] prothorax with strong keel and pinch, line = 0.40 mm; g.) ventral and h.) lateral view of Ischyrus n. sp. 7 [Panama] prothorax with prosternal keel swollen above pinch, line = 0.44 mm. 28 I use the term "prosternal plate" in reference to the surface of the prosternum between the coxal lines (Fig. 9). This "plate" is most often flat, but can be slightly convex. The plate shape varies throughout the genus (Fig. 11), from semicircular (wider than long) to elongate, parallel-sided (longer than wide). The length of the procoxal lines varies from species to species. I use three phrases to describe the length of these lines in relation to the procoxae: surpassing coxae, barely surpassing coxae, not surpassing coxae. The phrase "surpassing coxa" (Fig. lib) indicates the line passes beyond an imaginary line drawn between the anterior edge of the procoxae. The phrase "barely surpassing coxa" (Fig. lla, 11c) indicates the line passes beyond the point where the line in front of coxa begins, but does not pass beyond an imaginary line drawn between the anterior edge of the procoxae. The phrase "not surpassing coxa" (Fig. lid) indicates where the line stops where the line in front of procoxae begins, well before the imaginary line. Many species have sexual dimorphism on the prosternum in one or two forms. The majority of the species studied have a differing number, or varying development, of foveate punctures in front of the procoxa. In these species, females have more numerous or distinct foveate punctures than the males (Figs. 12-14), the opposite of Delkeskamp's (1959) observations of certain African Dacninae. a b c d Figure 11. Prosternal plates showing procoxal line shapes and anterior development, line = 1.0 mm; a.) Ischyrus proximus Lacordaire, barely suprassing coxa; b.) I. n. sp. 7, parallel-sided and surpassing coxa; c.) I. n. sp. 5, semicircular and barely surpassing coxa; d.) I. duponti Lacordaire, not surpassing coxa. Imaginary line (dashed) included for referencing anterior edge of procoxae in determining procoxal line development. Figure 12. Prosternal sexual dimorphism in puncture development. Male Ischyrus n. sp. 7 [Panama] prosternum: a.) ventral view, line = 0.38 mm; b.) lateral view, line = 0.50 mm; c.) ventral view of prosternal-pronotal epipleural suture, line = 0.17 mm; d.) prosternal punctures in front of procoxa, line = 0.04 mm. Female Ischyrus n. sp. 7 [Panama] prosternum; e.) ventral view, line = 0.43 mm; f.) lateral view, line = 0.50 mm; g.) prosternal punctures in front of procoxa, line = 0.04 mm. Figure 13. Prosternal sexual dimorphism in puncture development and lateral expansion. Male Ischyrus incertus Lacordaire [Mexico, Chiapas] prosternum: a.) ventral view, line = 0.75 mm; b.) lateral view, line = 0.50 mm; c.) ventral view of laterally expanded prosternal-pronotal epipleural suture, line = 0.17 mm. Female Ischyrus incertus Lacordaire [Panama] prosternum; d.) lateral view, line = 0.50 mm; e.) ventral view of prosternal-pronotal epipleural suture, line = 0.25 mm. Figure 14. Prosternal sexual dimorphism in puncture development and lateral expansion. Male Ischyrus scutellaris Gorham [Mexico, Yucatan] prosternum: a.) ventral view, line = 0.60 mm; b.) ventral view, line = 0.30 mm; c.) lateral view of prosternal expansion, line = 0.12 mm. Female Ischyrus scutellaris Gorham [Mexico, Yucatan] prosternum; d.) ventral view, line = 0.50 mm; e.) ventral view of prosternal-pronotal epipleural suture, line = 0.17 mm. 33 In a few species, males have the prosternum expanded laterally, obscuring the prosternal-pronotal epipleural suture (Figs. 13-14). The extent of the expansion appears consistent in all specimens of a species, but is variable among species. It can be a small expansion (Fig. 13), or it can nearly cover the entire pronotal epipleuron (Fig. 14). Mesosternum The mesosternum has few useful characters. The length of the mesosternal lines in relation to the distance between them, along with the meso-metasternal suture shape, is useful in grouping related species. This suture can be truncate (Fig. 15a), sinuate (Fig. 15f), or broadly sinuate (Fig. 15c). Metasternum The metacoxal lines usually extend posteriorly away from the medial side of the mesocoxa towards the hind angle of the metasternum (Fig. 9, 15). These lines are variable in shape and length. Anteriorly, the metacoxal lines can stop near the mesocoxa or continue along the meso-metasternal suture, often meeting at the middle. The shape of the line between the coxae varies from species to species and is useful in grouping related species. The term recurvedd" is used to describe a line that curves away from the meso-metasternal suture (Fig. 15c-f). Medially, these lines can take several forms: absent, not recurved nor meeting medially NMs d e f Figure 15. Meso- and metasternum showing coxal line development, line = 0.5 mm: a.) Ischyrus auriculatus Lacordaire, not recurved or meeting at middle; b.) I. aleator Boyle, not recurved and meeting at middle; c.) I. undulatus Gorham, recurved and meeting at middle; d.) I. n. sp. 13; e.) I. distinguendus Lacordaire, Mc = mesocoxa, Ms = mesosternum, Mt = metasternum; f.) I. n. sp. 3. 35 (Fig. 15a); not recurved meeting in a straight line (Fig. 15b); meeting with a series of punctures or undulations (Fig. 15c); meeting as a single tooth (Fig. 15e); meeting with two widespread teeth (Fig. 15d); or meeting with many teeth (Fig. 15f). The mesosternal line behind the mesocoxa is variable in its structure from species to species. It can be a simple line (Fig. 16a) which is single-sided, a groove (Fig. 16b) which is double-sided, or a groove which is notably deeper on one end and leads into a large pit (Fig. 16c). This pit is most often present at the medial end of this line, but the groove can be deepened at the lateral end (for example, I. n. sp. 13). First Visible Abdominal Sternite The coxal lines on the first visible abdominal sternite extend posteriorly from the medial side of the metacoxa. They are variable in length from specimen to specimen and are of little use in determining species. This first visible abdominal sternite can be rounded, broadly rounded or truncate between the metacoxa at the junction with the metasternum. Male Genitalia The internal sac of the male genitalia is held inverted within the median lobe (Fig. 17a). During copulation, the internal sac is everted, exposing any microstructure and extending the flagellum (Fig. 17b). The median lobe, internal sac, and flagellum are the true 9 Figure 16. Metasternal line posterior to mesocoxa: a.) simple line on Ischyrus proximus Lacordaire [Mexico, Chiapas], line = 0.75 mm; b.) groove on Ischyrus q. quadripunctatus (Olivier) [USA, Florida] line = 0.38 mm; c.) groove with pit at medial end on Ischyrus n. sp. 3 [Panama], line = 0.28 mm. Figure 17. Male genitalia of Ischyrus q. quadripunctatus (Olivier), line = 0.66 mm; a.) internal sac inverted as held within the body; b.) internal sac and flagellum everted as during copulation; F = flagellum, Is = internal sac, Ma = sclerite for muscle attachment at anterior end of flagellum, Ml = medial lobe, Ms = median strut. intromittent organs and show the majority of species specific characters. For additional insight into the genitalia of Coleoptera, I recommend the following: Lindroth 1957; Sharp and Muir 1912; Skelley 1993; Snodgrass 1957; Tuxen 1970; Verhoeff 1895; Williams 1945; and Wood 1952. The median lobe is a simple tubular structure that is curved and laterally flattened. The degree of curvature varies from species to species and can be slightly curved to arched. The shape of the median lobe's posterior end varies, and can be truncate, rounded, or narrowed and then rounded. The internal sac occasionally has pigmented areas, which are patches of microspinules. These can be large lightly pigmented patches of widely scattered microspinules, or small dark paired patches. These patches are known in only a few species. I found the shape of the sclerite for muscle attachment at the flagellum's base to be important in species recognition. The sclerites are all basically U- shaped, but with many variations. Just posterior to this sclerite is a pigmented section. This section of the flagellum is often variable in shape and curvature. A close study showed it to be flexible, like cartilage. The shape of this structure was not used to distinguish species because it can vary from specimen to specimen. The flagellum is variable throughout the genus: long, 39 cylindrical, and hair-like; flattened and ribbon-like; or straight and rigid. The tip of the flagellum can be pointed or flared. Female Genitalia Female genitalia (Fig. 18) varied little from species to species. Structures had proportions which varied, but species recognition based on female genitalia was not possible with any degree of confidence. The sclerotized spermatheca showed some variation in shape which could be useful in studies of higher categories. The spermatheca consisted of two parts; the head and tail. The head is the large bulbous, terminal structure, which varies in shape from circular to kidney-shape. Many species have spermathecae with a top-knot (bump on the head), variable in size and occasionally in position. The spermatheca tail is a sclerotized section of the duct extending from the spermatheca's head to the genitalia. The shape and thickness of the tail is variable, but similar in related species. Materials and Methods Specimens Dry preserved specimens were borrowed from many sources (institutions and individuals) during this revision. Appendix B lists these sources with their coden (mostly from Arnett et al., 1993) used throughout this study. S8- Tk H T a T8 aI Figure 18. Female genitalia of Ischyrus q. quadripunctatus (Olivier), line = 1.0 mm: a.) spermatheca; b.) ventral view; c.) dorsal view; A9 = abdominal segment IX, H = head of spermatheca, P1 = proctigeral lobe, S8 = abdominal sternum VIII, sS8 = straps appendant to abdominal sternum VIII, sT8 = straps appendant to abdominal tergum VIII, St = styli, T8 = abdominal tergum VIII, T = tail of spermatheca, Tk = top-knot on spermathecal head. Locating Types In most taxonomic research the investigators must study material seen by the describer of a species in creating the description. These specimens are the reference point for the name. Many descriptions were poorly done, or lack specific details needed to identify additional specimens. The history of each "type" specimen becomes important in locating it or discovering if it still exists. Modern researchers must record the depository of specimens studied. Early researchers did not always do this. Horn and Kahle (1935-1937) and Sachtleben (1961) stated the location or fate of many collections and can be helpful in locating a specific specimen. For New World Erotylidae the majority of early type material can be found in three places: the Museum National d'Histoire Naturelle, Paris (MNHN); the Crotch Erotylidae Collection, University of Cambridge, U.K. (CUMZ); and the Natural History Museum, London (NHML). Many of the nomina nuda in Dejean's catalogs (1836, 1837) and collection, were validated by Lacordaire because he acquired Dejean's Erotylidae (Horn & Kahle 1935-1937) and used Dejean's names in his descriptions. Horn & Kahle indicate that Lacordaire's collection was divided and deposited in various European museums. I have been unable to find all of Lacordaire's Erotylidae, but he studied specimens of several other collectors whose material can be 42 found in two places: the Crotch collection (CUMZ); or the Oberthir collection (MNHN). Many of Lacordaire's species were described from the Dupont collection. Horn & Kahle indicate that the Dupont collection was divided with some of the material being deposited in the collections of G. V. Mniszech and R. Oberthir. Nicole Berti at the Museum National d'Histoire Naturelle, Paris (in litt.), stated that the Oberthur collection, including the Mniszech collection, is at the MNHN, and that Oberthir indicated on the back of a box that specimens studied by Lacordaire are in the collection of Mniszech. For Ischyrus, the specimens labeled "Type" from this collection matched the original descriptions in both morphology and label data. Most specimens in this collection do not have "Type" labels, but they are potentially type material and should be considered in subsequent type designations for Lacordaire species. Lacordaire also studied specimens from the collections of L. A. A. Chevrolat and L. J. Rieche. These erotylid collections, and others, were acquired by G. R. Crotch. Crotch's Erotylidae collection, at the Cambridge University Museum of Zoology, is rich in types. In this collection, label data indicate which specimens are types, and often from whose collection they came. The Ischyrus specimens labeled "Type" fit the original descriptions, label data, and collection of origin. 43 The Natural History Museum, London, (formerly known as the British Museum of Natural History) houses the specimens studied in the Biologia Centrali-Americana (Gorham 1887- 1899). Tracking one specimen illustrates that it is not always a simple matter. The type specimen of Ischyrus quadripunctatus (Olivier), the generotype of Ischyrus, was not studied by Boyle (1956) in his family revision of America north of Mexico. Olivier (1791) stated that the specimen was in the collection of M. Francillon. Horn and Kahle (1935-1937) state that the Francillon collection was divided and deposited in the Natural History Museum, London, and the Hope Entomological Collections, University of Oxford, U.K. I visited the Natural History Museum and found no specimen which could be the type. The curator of the Hope Entomological Collections, G. C. McGavin, wrote that the majority of their part of the Francillon collection was sent to the (Alexander) MacLeay Museum, Sydney, Australia, in 1818. The curator of the MacLeay Museum, D. S. Horning, Jr., wrote that they do have specimens from the Francillon collection, including some from the locality stated in Olivier's description. But, he would be unable to look for the specimen until his recent injury had healed. Thus, I still do not know if the specimen studied by Olivier exists. Collecting Specimens of the genus Ischyrus can be found feeding on their host fungus. Based on the number of specimens known for most species, we still do not know where to look. Most specimens appear to have been haphazardly collected, even those collected in light traps. No definitive statement can be made about where to look for, or how to collect, members of this genus. I have collected various species of Ischyrus and related genera by using a beating square on dead sticks at night. Sticks, limbs, branches, etc., which produced the most specimens were either still on the tree, or on the ground, but suspended above the surface. The only known hosts are prostrate white fungi which seem to prefer dead suspended branches. Much work remains to be done in understanding the biology of this genus and the family. (See the Biology section under the Generic Account for additional comments.) Equipment Specimens were studied under a binocular dissection microscope, Unitron ZSB, with a zoom-magnification of 0.7x- 4.5x and 20x ocular lenses. A Hitachi S-570 scanning electron microscope was used. Although useful in seeing and understanding many of the minute characters, it was not an essential part of this study. I made an effort to base species descriptions and key characters on those visible at low magnifications. Techniques Adult specimens often needed to be cleaned before surface structures could be studied. This was accomplished by brushing the specimen with a small soft-bristle brush dipped in ethyl acetate or alcohol. Ethyl acetate was also used to degrease badly soiled specimens. If the mouthparts were badly soiled, the specimen was dipped in warm water to loosen the dirt, and to relax the specimen before brushing the dirt away. On rare occasions specimens were so badly soiled that they were totally relaxed, cleaned, and remounted. European-style card mounting made it impossible to study the ventral surfaces without removing the specimen from the card. The solvent used to soften the glue depended on what glue was used. The following series of chemicals was used until one dissolved the glue: water, 70% isopropanol, 80% ethanol; ethyl acetate. Rarely did the specimen require treatment with all of these, but occasionally a specimen needed to have the glue manually removed. I dissected many genitalia for further study. Specimens were chosen from across the distributional range of the species and from those showing variation in external characters. Each specimen was put into a weak solution of hot detergent water and allowed to sit from one hour to overnight. Once relaxed, the specimen was carefully held between the thumb and forefinger under the dissecting microscope. With a pair of jewelers forceps, the elytra were lifted just enough to slip one side of the forceps underneath and the abdomen was grasped on a side. In this way, the specimen was held and the remainder of the body was not effected by the dissection to follow. The abdominal tergites (membranous) and underlying muscle masses were separated from the visible sternites on the side not held by the forceps. This was done with a bent-tipped minute attached to a small wooden toothpick. Once loosened, the forceps were moved to hold only the sternites of the side just separated, and the same operation performed on the second side. After these separations were complete the tergites and underlying muscle masses were removed with a second pair of forceps. This technique allows the visible abdominal sternites to remain attached; the specimen appears intact. The removed muscle masses containing the genitalia, and terminal segments of the abdomen, were cleaned and cleared in a warm 10% potassium hydroxide (KOH) solution, and the remaining unwanted tissues were removed manually with forceps. The genitalia were rinsed in 70% isopropanol or water and stored in glycerin in genitalia vials associated with the appropriate specimen. Genitalia vials are small plastic or glass vials that can be placed under the pinned specimen, with the pin piercing the stopper. Detailed study of these genitalia rarely required more magnification than the dissecting microscope allowed. The 47 genitalia were studied in glycerin, and moved into various positions to see the shape of the muscle attachment at the anterior end of the internal sac. The shapes of this structure and the flagellum were important in solving many of the species problems. When finished, the genitalia were returned to the genitalia vial and pinned under the appropriate specimen. The stridulatoryy files" were not studied for many species since this required removing the head from the body. This type of dissection destroyed the specimen, and I was not willing to sacrifice the few specimens available for most species. Illustrations were made with the use of a glass-grid insert placed in an ocular of the dissecting microscope. The imposed grid on the specimen was used as reference and the drawing was made on a piece of grid paper. It was necessary to make various adjustment to each of the drawings, because the curvature of the lenses produced distortions. This was most apparent when comparing the finished pencil drawing and the specimen without the aid of the microscope. The drawings were then traced in India ink onto tracing paper with a 000 Koh-i-nor Rapidograph technical pen, scanned into electronic form using a Hewlett Packard Scanjet IIP, finished using CorelDRAW* 2.01 (a computer graphics program) on a Unisys 486 personal computer, and printed with an Apple LaserWriter" II. Habitus drawings lack legs and antennae for several reasons. First, the legs are of little use in determining species. The antennae need to be illustrated side-by-side for close comparisons. Adding these structures doubles or triples the time to do an illustration. Lastly, by omitting these structures, the habitus drawings can be placed closer together, requiring less space and fewer pages. In plotting distribution maps, many specific localities were not found because of poor label data. If the information was adequate enough find the general area, an open symbol was placed on the map in the general area. Specific localities are shown with solid symbols. If label data were vague and there was already a plotted locality in the general area, an additional symbol was not added. Questionable records are plotted with a question mark "?". Color Pattern Problems Species level decisions have historically been based simply on color and color patterns. Museum specimens vary in the shades of orange because of age, killing agent, and preservation technique. Because of this, the exact color is of little use in species determination. The color pattern is most important in determining species, but care must be taken in analyzing differences. The specimen(s) in question must first be placed within the proper section of the genus. This was done by using morphological and genitalic characters. Many recently 49 described species were compared to an unrelated species in the original description. This made determinations based on the literature impossible, and the type specimens had to be studied. A different appearing color pattern did not necessarily mean the specimen was a different species. For example, a species may have two spots on each elytron. When these spots are enlarged, they blend together and form a band. Differences based on changes in pattern due to the spot size generally were not specific. In other cases, a spot which varied in its location, generally indicated a specific difference. For example, in two closely related species the only color pattern difference is that one species has a humeral spot touching the base and the other has a subhumeral spot well removed from the base. In I. scriptus, I. proximus, I. palliatus, and I. incertus, the relative position of the pronotal spots and the shape of the circle they form is useful in determining species. Written descriptions cannot convey the exact details of these patterns the way an illustration does. Every species and many variations are here illustrated. In some cases the differences are subtle, but they are constant and correlate with other morphological differences. Care should be taken when comparing any specimen to these illustrations because of color pattern variations mentioned above, and ones not yet known. 50 Studying series of specimens from a large geographic range has allowed me to observe geographic variations in color patterns that were once considered specific. One such character is the color of the head, which can be red, black, or with some variation of both. In only a few cases has this variation been more than a clinal or subspecific difference. The term "pattern" was used for variations of a color pattern in situations where previously described species were found to be part of a dine. These "patterns" were maintained because they still had some relation to a geographic range. The "pattern" name is the specific name that once applied to that pattern, or a name was applied if that "pattern" was not previously described. New "pattern" names were applied only if specific names already existed for other patterns (see I. quadripunctatus and I. scriptus). These named "patterns" have no nomenclatural status. Another general trend is the change of elytral patterns from north to south. Many species have solid bands with smooth edges in the north; moving south, these edges become more and more sinuate. Some even develop into stripes as the sinuate edges on both sides of the band meet. Many species from mid-South America have striped patterns. In contrast, the majority of northern species have banded patterns. This is best illustrated with I. 51 quadripunctatus, I. scriptus, and their variations. This trend may indicate mimetic relationships among species. Rules of Thumb In determining the taxonomic status of various names, and in naming new taxa, several general rules were followed which require some explanation. Many species are variable in color pattern over their geographic range. This is illustrated in several species where adequate series have been studied. Certain color patterns are known from only a few specimens, often from scattered localities. If two specimens have different, but basically similar color patterns and their morphology (including the genitalia) is similar, they are considered variations of a single species. In other cases a radical color pattern difference is observed, but the genitalia are incomparable (i.e., male vs. female). These are considered variations of a single species, and are discussed under the most closely related described species, noting the variations and their taxonomic status. If a previously described species was a member of several patterns in a dine, that name was synonymized under the senior name and discussed in the species account. Consistent morphological differences correlated with color pattern differences are considered to be specific, and these taxa are described. New species are simply 52 numbered, because of problems with nomeclatural priority, and names will be proposed when published. The lack of series in many species indicates that there is much to be discovered in this genus. Several of the species discussed here may actually represent complexes. Because of inadequate material these problem taxa are left unresolved and are discussed under the appropriate species account. Results The genus Ischyrus is composed of 42 North and Central American species, including 16 previously undescribed; leaving 30 additional described species in South America. A total of 3741 North and Central American specimens was studied (2890 I. quadripunctatus) and 363 were dissected. The 148 figures, key, appendices, and descriptions are provided to complete this revision. ISCHYRUS LACORDAIRE Ischyrus Lacordaire 1842:89-131. Micrischyrus Alvarenga 1965:86. Type Species. Erotylus quadripunctatus Olivier 1791:431,437. Subsequent designation by Crotch 1873a:353; 1873b:144. Diagnosis. Characterized by having coarsely faceted eyes, triangular mentum, short ocular stria not surpassing anterior angle of eye, undilated tibia, and semicircular or trapezoidal antennomere IX. Description. Length 3.5 9.9 mm. Body shape parallel-sided, to elongate, or ovoid, slightly flattened to convex dorsally; microreticulation, surface dull to shining; unicolorous brown to variously banded or spotted, yellow- orange with black pattern. Head with ocular striae generally ending at or before anterior angle of eye, rarely extending onto epistome at base of antenna; frons often with an impression at each side near base of antennae; epistome wedge-shaped, generally with truncate apex; epistome punctures generally denser than punctures on vertex. Eye large, bulging from side; facets coarse (Fig. la), varying in size throughout the genus, rarely fine. 54 Antenna surpassing middle of pronotum, often reaching basal 0.25; antennomere I large, elongate; antennomere II circular, ball-like, length = 0.5 x antennomere I; antennomere III elongate, length equal to next 2 to 4 segments combined; antennomeres IV to VIII length subequal to width, length rarely more than 1.5 x width; antennomeres IV to VII rounded at ends; antennomere VIII edged and angled apically; antennomeres IX to XI form a loose club; antennomeres IX to X 3 to 4 x wider and 1.5 to 2 x longer than antennomere VIII; antennomere IX semicircular to trapezoidal, rarely triangular (Fig. 6); antennomere X crescent-shaped to trapezoidal; antennomere XI transversely elongate-oval to circular; antennomeres X-XI often asymmetrical. Mandibles each with two finger-like teeth and a large prostheca bearing many inwardly pointing setae. Maxilla with lacinia bearing an apical tooth, often bifid (Fig. 19b); terminal segment of palp triangular or securiform, width = 1 to 3 x length. Labial palpi vary from squared or circular to securiform, width = 1 to 2 x length (Fig. 7). Mentum with a pore on each side in front of basal corner; mental plate triangular, rarely longer than wide; ridge surrounding plate often raised laterally giving mentum a three prong crown-shape, medial prong (medial ridge extension) variously shaped, protruding or not (Figs. 8, 19a). Postmandibular lobes present, broadly rounded, Mp T --\ Lp o M a b Figure 19. Ischyrus q. quadripunctatus (Olivier) a.) labium and b.) left maxilla ventral view, line = 0.25 mm: M = mentum, Lp = labial palp, Mp = maxillary palp, T = bifid tooth of lacinia. Figure 20. Ventral view Ischyrus q. quadripunctatus (Olivier) [USA, Florida] head and prosternum, line = 0.50 mm. 57 forming inner side of groove next to the eye for reception of antennomeres II to III (Fig. 20). Pronotal disc evenly rounded; sides variably arched inwardly toward eyes; anterior angles closer together than posterior angles; anterior edge not margined between eyes; anterior angles forwardly produced, making anterior edge concave; base sinuate, not margined, lobed at middle, with group of large punctures at each side. Scutellum pentagonal, wider than long. Elytra with sides parabolically rounded to apex; 7 to 9 stria evident by rows of punctures, lacking at humerus and extreme apex, rarely impressed or missing; intervals flattened, often with minute punctures; base rarely margined; elytral epipleuron widest at base, strongly narrowed at hind coxae, gradually folding under to apex (Fig. 9); some elytral punctures each with a small protruding seta, visible in profile. Prosternum usually keeled, margined and constricted (pinched) at front (Figs. 10, 20); sternal plate shape and proportions variable (Fig. 11); lines anteriorly converging or parallel, rarely surpassing front of procoxa, lines not continuous around coxae (except n. sp. 1); posteriorly prosternum truncate or slightly concave, not margined. Mesosternal lines parallel or anteriorly divergent, straight or arched; plate square or transversely rectangular; posteriorly sinuate or truncate. Metasternal lines extending onto disc from inside of mesocoxa toward posterior angle of metasternum, rarely continuous around mesocoxae; variable in length, up to 0.5 distance to posterior angle; line behind mesocoxae variably impressed or grooved, occasionally with pit (Fig. 16). Legs with femora slightly swollen, complete margin on inner surface (Fig. 21); tibia straight, almost parallel- sided, slightly widened toward apex; tarsi pseudotetramerous. Abdominal coxal lines present, short; rarely continuous around metacoxae. Male genitalia with median strut length variable, equal to or larger than median lobe; internal sac can bear patches of spinules; flagellum varying in length and thickness, with sclerotized muscle attachment at base, non-slerotized section at base flexible; lateral lobes of tegmen generally flattened. Female genitalia with straps appendant to abdominal segment VIII; abdominal segment IX elastic, length variable; flattened plate-like proctigeral lobe; apical segment of coxite with slender styli (Fig. 18). Proportions of these stuctures vary little throughout the genus. Spermatheca sclerotized, shape of head and tail variable, occasionally with a top-knot (Fig. 18). Stridulatory files often present at the base of the head (Fig. 22). Figure 21. Ventral view of meso-femur: a.) Ischyrus q. quadripunctatus (Olivier) [USA, Florida] with posterior margin, line = 0.50 mm; b.) Oocyanus flavitarsis (Lacordaire) [Cuba] lacking posterior margin, line = 0.60 mm. * q.'~S Figure 22. Ischyrus q. quadripunctatus (Olivier) [USA, Florida] occipital region of head showing stridulatory file (arrow): male with stridulatory file, a.) line = 0.38 mm, b.) line = 0.08 mm; c.) female lacking stridulatory file, line = 0.15 mm. 61 Sexual dimorphism often present. Males of some species have the prosternum laterally expanded onto the pronotal epipleuron; not expanded in females (Figs. 13-14). Males of many species have fewer, or less distinct, punctures on the prosternum in front of the procoxae (Figs. 12-14). Distribution. This genus is restricted to the New World, where it is widespread, occurring from southeastern Canada near the St. Lawrence Seaway and southeastern North Dakota through the eastern U.S. and southern Arizona, Mexico, Central America, the West Indies, into South America to northern Argentina. Bioloav. The life history of this genus is basically unknown. Only one species has its larva described; the type species I. q. quadripunctatus (Olivier) (see Weiss 1920, Skelley 1988b, Chapuis & Candeze 1855, Chapuis 1876). The following is based on published accounts of this species, a few bits of information taken from label data, and personal communications and observations. The larva of I. q. quadripunctatus has well developed dorsal sclerotization with short spines. It is unusual in that the pronotal sclerotized area is broken into parts appearing like false eyes (Fig. 23). Members of a related genus, Oocyanus Hope, have a similar set of "eye-spots". This peculiarity could be of adaptive significance in warding off predators. Both of these larvae have been found feeding exposed on prostrate white fungus growing on dead wood (personal observations). Other larvae of the Figure 23. Ischyrus q. quadripunctatus (Olivier) larva [USA, Florida], line = 4.0 mm.; a.) lateral view; b.) dorsal view of head and thorax. _ _ Erotylidae are burrowers in fungi (some Triplacinae, Dacninae) or are surface feeders (some Erotylinae). The surface feeding Erotylinae are often protected by a covering of large spines and often have color patterns; whereas burrowing species lack these spines and patterns (see Roberts 1958; Costa, et al. 1988; and Lawrence 1991, for illustrations of various erotylid larvae). Ischyrus q. quadripunctatus has been collected on a white resupinate polypore fungus, Oxyporus latemarginatus (Dur. & Mont. ex. Mont.) Donk, also known as Poria ambigua Bres. (Skelley, et al. 1991). Richard Leschen (pers. comm.) collected I. proximus on Schizopora paradoxa (Fr.) Donk in Costa Rica, also a white resupinate polypore. Both of these fungi are white rot fungi of wood. Adults have been taken by general collecting methods; in leaf litter, under bark, sweeping vegetation, etc. Many specimens have been taken at light, suggesting nocturnal activity. This is also indicated by the large eye facets present in members of this genus. Using a beating square at night, I collected several species of Ischyrus and other related genera on small dead limbs both on the ground and hanging from the trees. Several species have pits on one of the thoracic sternites, for example; I. undulatus and I. n. sp. 3 on the metasternum behind the mesocoxa, and I. n. sp. 1 on the prosternum. These pits occur on both sexes of the species and show no sexual dimorphism in their development. Their 64 function is not known, but they could be used for structural support, muscle attachment, areas for glandular secretion, or mycangia (Crowson 1981). Etymoloav. Ischyros: Greek for strong, mighty, excessive (Brown 1985). Possibly named in reference to the strongly clubbed antennae or pronounced color patterns. Gemminger and Harold (1876) indicated that the name Ischyrus means "validus". Remarks. Although Boyle (1956:110) stated that Ischyrus lacks teeth on the lacinia; this is incorrect. The tooth illustrated (Fig. 19) is located in a dense patch of setae and is difficult to see. Ischyrus Lacordaire appears most closely related to Megischyrus Crotch, Callischyrus Crotch, and Oocyanus Hope in having a triangular mentum, loose antennal club, strongly microreticulate body surface, and in basic color patterns. Ischyrus differs from Oocyanus in having the femora margined along the inner side where the tibiae meet the femora (Fig. 21a); this margin is lacking in Oocyanus (Fig. 21b). Callischyrus differs from Ischyrus in having the eyes finely faceted and the ocular stria surpassing the antennal base; in Ischyrus the eyes are coarsely faceted and the ocular stria at most touch the antennal base. Megischyrus differs from Ischyrus in having a larger body size (greater than 11 mm) and in having antennomere IX triangular (Figs. 6n-o); in Ischyrus the body size is smaller (less than 10 mm) and antennomere IX is semicircular or trapezoidal, rarely triangular. References. Alvarenga 1965:85; Arnett 1963:817-821; 1985:341-342; Blackwelder 1945:465; Boyle 1956:132-137,128; Chapuis 1876:35-38; Crotch 1873a:353-354; 1873b:144; 1876:426-433(50-57); Curran 1944:1-5; Edwards 1949:94; Gemminger & Harold 1876:3690-3691; Germar 1843:133; Girard 1873:820; Gorham 1887:39-45; Kuhnt 1909:55,57,61-64; 1911:42-44; Lacordaire 1842:89-131; LeConte & Horn 1883:124; Leschen 1991:180, 192; Mader 1942:171,195-196; 1951:209-210; Neave 1939-1940:790; Pallister 1955a:4; 1955b:6-7; Seidlitz 1891:288; Skelley 1988b:60. ARTIFICIAL KEY TO SPECIES This key was built using characters visible under low magnification, without dissection. Several species appear in the key more than once, beacuse some characters are variable or have an intermediate state on some species. Couplet 3 is the best example of a character that can be difficult to interpret. If a specimens does not adequately key, or does not match what it does key to, then try the other choice in the couplet. 1. Antennomere IX triangular, sides straight, as long or longer than wide (Figs. 6k, 6n-o); body convex dorsally (Fig. 2b) ........................................... 2 1'. Antennomere IX trapezoidal to semicircular, sides angled or rounded, generally wider than long (Figs. 6a-j, 61- m); body parallel-sided, elongate or ovoid; flattened above (Fig. 2) ...................................... 3 2.(1) Pronotum with 2 free spots (Fig. 24). . .............................. vespertilio Lacordaire 2'. Pronotum with central stripe (Fig. 25). ..... n. sp. 1 3.(1') Antennal club segments distinctly asymmetrical; antennomere XI larger (wider or longer) than antennomere X (Fig. 6a-f) ........................... 4 3'. Antennal club segments symmetrical; antennomere XI size variable; if appearing asymmetrical, antennomere XI 66 24L 25 26 27 I 1 28 29 30 31 32 33 34 35 Figures 24-35. Dorsal habitus, line = 1.0 mm: 24.) Ischyrus vespertilio Lacordaire; 25.) I. n. sp. 1; 26.) I. n. sp. 2; 27.) I. distinguendus Lacordaire; 28.) I. insolens Crotch; 29.) I. scriptus (Olivier) "northern"; 30-31.) I. scriptus "southern"; 32.) I. bogotae Crotch; 33.) I. incertus Lacordaire; 34.) I. proximus Lacordaire; 35.) I. angularis Lacordaire. 68 equal in size or smaller than antennomere X (Fig. 6g- m). .............................................. 19 4.(3) Pronotum without free spots ........................ 5 4'. Pronotum with free spots ............................. 6 5.(4) Pronotum with a central black stripe and red sides, no basal spots (Fig. 26). ................... n. sp. 2 5'. Pronotum red with three basal spots only (Fig. 27). . ............................ distinguendus Lacordaire 6.(4') Pronotal hind angles with a dark marking, black of lateral margin encroaches upon the side ............. 7 6'. Pronotal hind angles without distinct dark markings .. 8 7.(6) Pronotum with 4 free spots in a transverse arc; 2 central spots well separated from the base (Fig. 28). . ..................................... insolens Crotch 7'. Pronotum with 2 basal, 2 central, and 2 anterior spots forming a circle; 2 central spots close to or connected with the base (Figs. 29-31) ...... scriptus (Olivier) 8.(6') Pronotum with 2 free spots; 2 basal spots rarely free or weakly touching margin; if appearing free, then spots separated from base by less than their diameter (Figs. 29, 32-35). .................................. 9 8'. Pronotum with 3-4 free spots; if 4, then central spots separated from base by more than their diameter (Figs. 37-47). .. .......................................... 14 9.(8) Pronotum with spots on anterior margin. ......... 10 9'. Pronotum without spots on anterior margin ......... 13 69 7-, (At-\,|I \ 7, 0;; * 46 Figures 36-47. Dorsal habitus, line = 1.0 mm: 36.) Ischyrus septemsignatus Gorham; 37.) I. tripunctatus Crotch; 38.) I. frontalis Lacordaire; 39.) I. frontalis var.; 40.) I. dunedinensis Blatchley; 41.) I. boucardi Crotch; 42.) I. quadripunctatus chiasticus Boyle; 43- 47.) I. quadripunctatus quadripunctatus (Olivier); 43.) I. q. q. "quadripunctatus"; 44.) I. q. q. graphicsus; 45.) I. q. q. "subcyindricus"; 46.) I. q. q. "Antillean"; 47.) I. q. q. "banded-leg". \7I ** (j 1;& \% 44 10.(9) Pronotum with 1 large basal spot (Fig. 32). . ...................................... bogotae Crotch 10'. Pronotum with 2 basal spots ........................ 11 11.(10') Antennomere XI equal in size to antennomere X (Fig. 61); femur black; elytra with weak microreticulation, strongly shining; pronotal spots forming a transversely elongate ellipse (Fig. 33) ........ incertus Lacordaire 11'. Antennomere XI larger than antennomere X (Fig. 6c); femur red or black; elytra with dulling microreticulations; pronotal spots forming a circle, longitudinally elongate circle, or rarely a weak transverse ellipse (Figs. 29-31, 34) ............... 12 12.(11') Femur red with dark knee; humeral spot free from scutellar spot; elytral epipleural fold black, occasionally pale in general specimens; pronotal spots form a circle or weakly transversely elongate ellipse (Figs. 29-31) ..................... scriptus (Olivier) 12'. Femur and scutellum black; humeral spot connected to scutellar spot; elytral epipleural fold red, at least at base, rarely dark; pronotal spots forming a longitudinally elongate ellipse (Fig. 34).............. ................................. proximus Lacordaire 13.(9') Elytra with central band and apical spots connected to suture (Fig. 35) .............. angularis Lacordaire 13'. Elytra with central band and apical spots not reaching the edge (Fig. 36) .............. septemsignatus Gorham 14.(8') Pronotum with 3 free spots ..................... 15 71 14'. Pronotum with 4 free spots ........ ................. 17 15.(14) Central elytral band broken into spots .......... 16 15'. Central elytral band complete (Fig. 37) . .. ............................ tripunctatus Crotch 16.(15) Lateral spot of elytra small, circular; Central America (Figs. 38-39). ......... frontalis Lacordaire 16'. Lateral spot of elytra elongate, stripe-like; USA, Florida (Fig. 40). dunedinensis Blatchley 17.(14') Maxillary palps narrow (Fig. 7f), longer than wide. ...................... quadripunctatus (Olivier). .18 17'. Maxillary palps wide (Fig. 7c), wider than long; (Fig. 41) ................................... boucardi Crotch 18.(17) Scutellar spots form an X-shaped mark; narrow band of black at base of pronotum (Fig. 42) . ................... quadripunctatus chiasticus Boyle 18'. Scutellar spots not X-shaped, or if X-shaped then base of pronotum with prominent tooth-like spots (Figs. 43- 47) ....... quadripunctatus quadripunctatus (Olivier) 19.(3') Metasternal line behind mesocoxa impressed with a distinct pit near middle (Figs. 15c, 15f, 16c); epistome angled at side, flat and sharp in profile; face flat, usually lacking impressions; body cylindrical ................. ...... ................. 20 19'. Metasternal line behind mesocoxa without pit, often impressed (Figs. 15a-b, 15d-e, 16a-b); epistome angled or not, often thickened at apex and rounded in profile; face often with impressions; body often flattened .. 21 20.(19) Pronotum with 2 basal spots and a central stripe connecting the base and anterior margin (Fig. 48). . ........................................ n. sp 3 20'. Pronotum without a central stripe, 4 free spots (Fig. 49). ................................. undulatus Gorham 21.(19') Color pattern lacking, uniformly colored, often paler at lateral margins. ........................ 22 21'. With distinct color pattern ........................ 25 22.(21) Body with metallic blue sheen; strial punctures present only at base and along sutural margin; Cuba (Fig. 50) ...................... ............. n. sp. 4 22'. Body without metallic sheen; strial punctures present over entire elytral disc ........................... 23 23.(22') Prosternum 2-3 times longer than the distance between the procoxae (Fig. 11b); labial palps not expanded, squared or rounded (Fig. 7k) ............. 24 23'. Prosternum at most 1.5 times longer than the distance between the procoxae; labial palps expanded, securiform (Fig. 7j); (Fig. 51) .......................... n. sp. 5 24.(23) Pronotal punctures distinctly larger laterally; shiny black; parabolically rounded at sides (Fig. 52) ............................................ n sp 6 24'. Pronotal punctures same size throughout; dull brown; parallel-sided (Fig. 53). .................... n. sp. 7 25.(21') Pronotum without free spots .................... 26 25'. Pronotum with free spots .......................... 32 56 57 58 59 Figures 48-59. Dorsal habitus, line = 1.0 mm: 48.) Ischyrus n. sp. 3; 49.) I. undulatus Gorham; 50.) I. n. sp. 4; 51.) I. n. sp. 5; 52.) I. n. sp. 6; 53.) I. n. sp. 7; 54.) I. aleator Boyle; 55.) I. n. sp. 8; 56.) I. auriculatus Lacordaire; 57.) I. auriculatus var.; 58.) I. ephippiatus Gorham; 59.) I. n. sp. 9. 26.(25) Prosternum not strongly constricted at anterior margin, not produced in profile, not pinched (Fig. 10a). ............................................ 27 26'. Prosternum strongly constricted at anterior margin, produced in profile, pinched (Fig. 10c-f). ......... 30 27.(26) Each elytron with 1 central stripe-like spot (Fig. 54) ..................................... aleator Boyle 27'. Elytra banded or spotted ........................... 28 28.(27') Pronotum entirely pale, elytra with small spots (Fig. 55) .................................... n. sp. 8 28'. Pronotum black or with black markings, elytral marking variable. ......................................... 29 29.(28') Elytra with distinct free spots, not banded (Fig. 25); metasternal coxal lines continuous behind mesocoxae; line behind mesocoxa not impressed; prosternal line in front of coxa a pit-like groove (Fig. 9b) .................................... n. sp. 1 29'. Elytra distinctly banded (Figs. 56-57); metasternal coxal lines not continuous behind mesocoxae, long; line behind mesocoxa impressed, groove-like; prosternal line in front of coxa simply impressed. . . .............................. auriculatus Lacordaire 30.(26') Body stout, rounded laterally; pronotum entirely black, or with anterior angles pale; each elytron with a free basal spot .................................. 31 30'. Body parallel-sided, elongate; pronotum with pale anterior angles and often with central red markings; 75 elytra lacking free basal spots (Fig. 58) . .................................. ephippiatus Gorham 31.(30) Free scutellar and humeral spots in transverse line, central elytral fascia not extended to apical quarter (Fig. 59) ............................... n. sp. 9 31'. Free humeral spot located behind free scutellar spot; central elytral fascia reaching apical quarter (Fig. 60) ......................................... n. sp 10 32.(25') Pronotum with 2 free spots and three basal spots (1 band-like free spot in collatinus), 3 basal spots not including possible posterior angle spots ........... 33 32'. Pronotum with 2 or more free spots, possibly with three basal spots, but not with the above combination .... 37 33.(32) Pronotal hind angle with a spot or widening in the black margin; often with 2 weak black spots at anterior margin of pronotum (Fig. 61) ................ n. sp. 11 33'. Pronotal hind angle without spot, lacking spots at pronotal anterior margin ........................... 34 34.(33') Two free pronotal spots circular; scutellar spot often broadly connected to elytral base ............ 35 34'. Free pronotal spot(s) transversely elongate, rectangular; scutellar spot not connected to elytral base ............................................... 36 35.(34) Apical elytral spot free; central band not reaching lateral margin (Figs. 62-63) ............ pictus Gorham 35'. Apical elytral spot broadly connected to suture and apex; central band complete, reaching lateral margin (Fig. 35) ................ angularis Lacordaire 76 60 61 62 63 S 64 65 66 67 68 69 70 71 Figures 60-71. Dorsal habitus, line = 1.0 mm: 60.) Ischyrus n. sp. 10; 61.) I. n. sp. 11; 62.) I. pictus Gorham; 63.) I. pictus var.; 64.) I. episcaphulinus Gorham; 65.) I. collatinus Crotch; 66.) I. elegantulus Lacordaire; 67.) I. elegantulus var.; 68.) I. chacojae Gorham; 69.) I. chacojae var.; 70.) I. scutellaris Gorham; 71.) I. scutellaris var. 36.(34') Pronotum with 2 free rectangular spots; lacking sutural spot near elytral apex (Fig. 64) . ............................... episcaphulinus Gorham 36'. Pronotum with 1 transverse spot; with a sutural spot near the elytral apex (Fig. 65) ..... collatinus Crotch 37.(32') Pronotum with 2 free spots, base at most with thin black margin, rarely with anterior spots ........... 38 37'. Pronotum variously marked with 2-4 discal spots, other markings variously connected to margins ............ 42 38.(37) Scutellar spot narrowly connected to elytral base at scutellum, connection narrower than the spots' width (Figs. 66-67). ................. elegantulus Lacordaire 38'. Scutellar spot broadly connected to elytral base, connection same width as spot ...................... 39 39.(38') Humeral and scutellar spots usually separated; scutellar spot broadly connected to suture; antennomere IX rounded or angled at base, not triangular ....... 40 39'. Humeral and scutellar spots connected, appearing as one, separated from suture; antennomere IX triangular (Fig. 24). ... .................. vespertilio Lacordaire 40.(39) Head entirely red; humeral spot not connected to elytral base ....................................... 41 40'. Head with at least base black; humeral spot connected to elytral base, often connected to scutellar spot (Figs. 68-69). ....................... chacojae Gorham 41.(40) Central elytral band broken into round spots (Figs. 70-71) ............................. scutellaris Gorham 41'. Central elytral band broken into longitudinally elongate spots (Fig. 72) .................... n. sp. 12 42.(37') Pronotum with 3 free spots ..................... 43 42'. Pronotum with 2 or 4 free spots .................... 45 43.(42) Central elytral fascia complete (Fig. 37) . ................................. tripunctatus Crotch 43'. Central elytral fascia broken into spots. ........ 44 44.(43') Lateral spot of elytra small, circular; Central America (Figs. 38-39) ............ frontalis Lacordaire 44'. Lateral spot of elytra elongate, stripe-like; USA, Florida (Fig. 40). ............. dunedinensis Blatchley 45.(42') Pronotum with 2 free, 2 basal, and 2 anterior spots (Fig. 33). ........................ incertus Lacordaire 45'. Pronotum with 4 free spots; or with pronotal disc markings based on 4 spots in a transverse line; 2 central spots occasionally connected to the anterior margin, giving it the appearance of 2 free spots ... 46 46.(45') Pronotum with 4 free spots only; occasionally with dark base, but no distinct basal spots ............. 47 46'. Pronotum with 4 free spots and additional markings connected to margins ................................ 48 47.(46) Elytral central spot not connected to suture (Fig. 73) .......................... ............... n. sp. 13 47'. Elytral central band complete, one continuous marking connected to suture, possibly reaching lateral margin (Fig. 74) ......................... tetrasticus Gorham (0a \ Figures 72-78. Dorsal habitus, line = 1.0 mm: 72.) Ischyrus n. sp. 12; 73.) I. n. sp. 13; 74.) I. tetrasticus Gorham; 75.) I. n. sp. 14; 76.) I. n. sp. 15; 77.) I. fulmineus Delkeskamp; 78.) I. n. sp. 16. eS 5 4 (001 di9 74 /0\ 48.(46') With distinct anterior pronotal markings, or markings connecting discal spots to anterior margin 49 48'. Without distinct anterior pronotal marks, discal spots free (Fig. 75) .............................. n. sp. 14 49.(48) Each pronotal hind angle with a spot, occasionally reduced and appearing as dark swelling at margin of disc ............................................... 50 49'. Pronotal hind angles without spots, lateral margin dark but not extending onto disc (Fig. 76) ....... n. sp. 15 50.(49) Humeral spot connected to elytral base; central elytral band divided by orange except at stria V & VI (Fig. 77). .................... fulmineus Delkeskamp 50'. Subhumeral spot not connected to elytral base; central elytral band completely divided by orange (Fig. 78). . ....................................... n. sp 16 SPECIES ACCOUNTS Ischyrus aleator Boyle Ischyrus aleator Boyle 1954:46-48. Diagnosis. Unique in Ischyrus by its linear elytral spots (from base at humerus to apical third), pronotum lacking free spots, and weakly pinched prosternum. Description. Length: 5.6-7.4 mm; Width: 2.5-3.4 mm. Body elongate, parallel-sided, widest at basal third elytra; strongly microreticulate, dull; pale orange with black pattern (Fig. 54). Head orange, often with black epistome. Pronotum entirely edged in black; anterior edge with 2 spots; basal edge with 2 spots farther apart than anterior spots; spots occasionally touching on disc. Scutellum black. Each elytron with black epipleural fold, often with pale base; suture finely edged in black; disc with elongate triangular stripe, widest anteriorly, narrowly connected at base near humerus, reaching apical quarter of elytra. Ventral color variable from mostly black to orange with black sclerite edges. Legs black, femur occasionally banded with pale orange. Head dorsal distance between eyes = 2.3 x eye width; ocular striae reaching antennal base; vertex and epistome puncture size = 1 x facet, separated on vertex by 2 diameters, separated on epistome by 1 diameter. Antenna reaching base of pronotum; antennomere III as long as next 3 antennomeres combined; antennomeres IX-XI symmetrical; antennomere XI oval, circular to transverse (similar to Fig. 6i). Maxillary palp terminal segment semicircular; medial edge straight at base, angle 90; lateral side rounded, angle obtuse; width = length. Labial palp terminal segment triangular, extended on medial side, narrow, sides rounded, width = 0.8 x length. Labial palp width = 0.5 x maxillary palp width (Fig. 7m). Mentum with plate broadly triangular, length = 0.5 x width, sides slightly convex, ridge medial extension acutely pointed (Fig. 8i) Pronotal disc puncture size = 1 x facet, separated by 2 to 3 diameters; lateral punctures slightly larger and denser, separated by 1 to 2 diameters. Scutellum pentagonal, length = 0.6 x width. Each elytron with 7 visible striae; strial puncture size = 0.5 x lateral pronotal puncture size, becoming finer toward apex; intervals finely punctate, obscured by strong microreticulation. Prosternum not keeled, convex; anterior pinch weak, if present (Fig. 10a-b); with (female) or without (male) foveate punctures in front of procoxa; coxal lines nearly straight, length = 0.5 x sternal length, lines not surpassing coxae, length = 0.6 x basal width; prosternal plate flat, apical width = 0.75 x basal width; base shallowly concave. Mesosternum basal width = 2 x mesocoxal line length; coxal lines straight, parallel to anteriorly divergent; base sinuate. Metasternum coxal lines meeting at middle in straight line with a row of punctures, often weak (Fig. 15b); coxal line length variable, continuous around coxae or reaching a maximum of 0.33 distance to posterior angle of metasternum; line behind mesocoxa deep, groove-like; sternum medial punctures fine, few coarse lateral punctures. First visible abdominal sternite with coxal lines reaching 0.25 to 0.5 distance to posterior margin; rounded between metacoxae. Male genitalia with median lobe weakly arched, narrowed and apically rounded; internal sac without noticeable sclerotized structures; flagellum long and narrow, straight and apparently rigid at basal 0.5, length = 2 x median lobe length (Fig. 79a); base of flagellum straight, sclerite at base elongate claw-shaped (Fig. 79b-d) (9 northern Mexican, 1 Central American dissected). Female genitalia with spermathecal head cone-shaped; tail swollen and recurved onto itself at middle (Fig. 79e) (6 northern Mexican dissected). Stridulatory files present on occipital region of males' heads; absent on females. Males with few or no foveate punctures on prosternum in front of procoxae; females with few to many punctures on prosternum in front of e79 d 9" d c od c d c d c a ---- - Figures 79-83. Genitalia: 79.) Ischyrus aleator Boyle [male, Mexico, Chihuahua; female, Mexico, Sonora]; 80.) I. angularis Lacordaire [male & female, Panama]; 81.) I. auriculatus Lacordaire [male, Mexico, Chiapas; female, Mexico, Veracruz]; 82.) I. bogotae Crotch [male, Costa Rica, Puntarenas; female, Ecuador, Pichincha]; 83.) I. boucardi Crotch [male, Panama; female Peru, Madre de Dios]; a.) male genitalia, line = 0.66 mm; b.) lateral view, c.) dorsal view, and d.) anterior view of the sclerotized muscle attachment at anterior end of male flagellum, line = 0.22 mm; e.) female spermatheca, line = 0.33 mm. 85 procoxae. There is some overlap in the numbers of visible punctures between males and females. Variation. Markings vary dramatically in the width on specimens from north to south. Pronotal markings often touch in the middle, becoming stripe-like, leaving 3 pale circles. The width of the elytral stripe varies from 1 to 3 strial intervals. Central American specimens have banded legs, an orange elytral epipleural fold, and black epistome. Specimens from Northern Mexico and Arizona have entirely black legs, black epipleural fold, and an orange epistome. The type specimen, which appears to be slightly general, has brown legs. Variation in mesocoxal line, from parallel to anteriorly divergent, did not correlate with geographic range. Specimens from Arizona and northern Mexico have the metacoxal line short, reaching a quarter of the distance to the posterior angle, or continuous around the coxae. Specimens from southern Mexico and Central America have the metacoxal lines longer, nearly reaching a third of the distance to the posterior angle. The prosternal pinch is not present on the holotype. On other specimens this pinch varies from absent to weak but distinct. Type. The holotype (original designation) of Ischyrus aleator Boyle label data: "/ Cave Creek, Cochise Co./ Chiricahua Mts., AZ., 7000ft, June 24, 1927/ J.A.Kusche Collector/ Van Dyke Collection/ [red] Holotype Ischyrus aleator Boyle/" [CASC, Type #8369, studied]. Sex male. Specimens examined. The holotype and 44 specimens, representing 20 collection records, were studied (see Appendix C for specific data). Distribution. Southeastern Arizona, Mexico, El Salvador, and Guatemala (Fig. 84). Etymology. aleator: Latin = dice layer, gambler. Name alludes to the resemblence of the trifoliate tawny spot on the pronotum to the emblem of the club suit in a deck of cards (Boyle 1954). Taxonomic notes. The variation in leg color, size of elytral stripe, and development of metacoxal lines, appear to be correlated with geographic range. Few specimens with banded legs were studied from central Mexico and Central America. The importance of this color variety is uncertain. References. Boyle 1956:133,136-137,169 f.131-132; Skelley et al. 1991:65. Ischyrus angularis Lacordaire Ischyrus angularis Lacordaire 1842:126 Ischyrus quinquepunctatus Gorham 1887:43-44, t.3 f.6, new synonymy. Diagnosis. Characterized by its elongate body, complete central elytral band, apical elytral spots touching margins, and 2 free and 3 basal pronotal spots. Description. Length: 5.0-5.8 mm; Width: 2.4-2.9 mm. Body elongate, parallel-sided, widest at basal third of ',~ p b P. Figure 84. Ischyrus aleator Boyle [circle] and I. angularis Lacordaire [star] distribution map. 3 'r ~ "r d' 88 elytra; weakly microreticulate, shining; yellow-orange with black pattern (Fig. 35). Head entirely black, or with orange epistome and frons. Pronotum with 2 free spots and 3 basal spots; medial basal spot small; lateral basal spots larger, tooth-like. Scutellum black. Each elytron with orange epipleural fold; elongate subhumeral spot connected to base, often narrowly connected to large scutellar spot at base; scutellar spot broadly connected to base and suture; suture finely edged black; central band connected to lateral margin and suture; apical spot broadly connected to suture and apical margin; lateral margin black from central band to apex. Venter black, except for the orange lateral abdominal sternites and pronotal epipleuron. Legs black, tarsi brown. Head dorsal distance between eyes = 2.2 x eye width; ocular striae reaching 0.8 to 1.0 distance to anterior angle of eye; vertex puncture size = 1 x facet, separated by 1 to 2 diameters; epistome puncture size = 0.5 x facet, separated by 1 diameter. Antenna reaching basal 0.25 of pronotum; antennomere III as long as next 3 antennomere combined; antennomeres X to XI asymmetrical; antennomere XI transverse (Fig. 6e). Maxillary palp terminal segment triangular, securiform, basally rounded, apical angles nearly 900, length = 0.75 x width. Labial palp terminal segment triangular, extended on medial side, rounded basally, length = 0.66 x width. Labial palp width = 0.75 x maxillary palp width (similar to Fig. 89 7b). Mentum with plate broadly triangular, length = 0.6 x width, sides straight, ridge medial extension acutely pointed (similar to Fig. 8d). Pronotal puncture size = 1 x facet, separated by 1 to 2 diameters; punctures smaller and denser at extreme lateral edge. Scutellum pentagonal, length = 0.75 x width. Each elytron with 7 complete striae; stria VIII weak, visible on apical half; strial puncture size = 2 x pronotal disc punctures, gradually decreasing in size posteriorly; intervals finely punctate. Prosternum keeled and pinched anteriorly; with (female) or without (male) foveate punctures in front of procoxa; coxal lines straight, length = 0.5 x sternal length, lines surpassing coxae, length = basal width; prosternal plate flat, apical width = 0.6 x basal width; base shallowly concave. Mesosternum basal width = 2.5 x mesocoxal line length; coxal lines straight; base sinuate, lobed medially. Metasternum coxal lines not meeting at middle; coxal lines extend 0.5 distance to posterior lateral angle; sternum with medial punctures fine, few coarse lateral punctures separated by 2 to 3 diameters. First visible abdominal sternite with coxal lines reaching 0.5 distance to posterior edge; rounded between metacoxae; coarse punctures laterally, fine punctures medially. 90 Male genitalia with median lobe weakly arched, apically truncate with constriction just before tip; internal sac with end near median lobe roughened; flagellum long and narrow, length = 1.6 x median lobe length (Fig. 80a); base of flagellum straight, sclerite at base claw-shaped (Fig. 80b-d) (2 dissected). Female genitalia with spermathecal head kidney-shaped, often with top-knot; tail swollen, weakly curved (Fig. 80e) (3 dissected). Presence of stridulatory files on occipital region of head unknown (heads retracted). Females have foveate punctures on prosternum in front of procoxae; males lack these punctures. Variation. One specimen with a red head, probably general, has the same collection data as a black headed specimen. Types. The lectotype (here designated) of Ischyrus angularis Lacordaire label data: "/ Colombie/ Angularis Lac./ Type/ Ex-Musaeo Mniszech/ [red] TYPE/ [pale blue] Museum Paris ex Coll. R. Oberthiir 1952/ [red] LECTOTYPE Ischyrus angularis Lacordaire des.P.E.Skelley 1993/" [MNHN, studied]. Sex apparently female, abdomen missing. The holotype (by monotypy) of Ischyrus quinquepunctatus Gorham label data: "/ [red circle on white paper] Type/ Type sp. figured/ Bugaba, Panama, Champion/ Ischyrus 5-punctatus Gorham/ B.C.A., Col.,VII, Ischyrus/" [NHML, studied]. Sex not determined. |