The Biogeography and Phylogeny of Hologymnetis (coleoptera: Scarabaeidae: Cetoniinae) with a Revision of the Genus

The Neotropical genus Hologymnetis (formerly Cineretis) is comprehensively reviewed. Descriptions and a key to the seven species are presented, including three new species (H. kinichahau from Guatemala and Mexico, H. moroni from Mexico, and H. vulcanorum from El Salvador). Hologymnetis margaritis is restored to species status. A cladistic analysis reconstructing the presumed phylogeny of the genus is provided, and a discussion of the characters and their states is given. Hologymnetis cinerea and H. undulata were found to be the most derived species, while H. vulcanorum, H. margaritis and H. kinzchahau are the least derived; H. moroni and H. argenteola are sister species. The biogeographical study indicates that the genus is South American in origin with one species (H. undulata) now occumng there south of the Amazon Basin. Dispersal through Central America after the formation of the Panamanian land bridge is indicated. Descendent taxa then dispersed and speciated from El Salvador through southern and western Mexico and extreme southern Arizona. Subsequent formation of unfavorable habitat (primarily lowland rainforest) from Amazonia to Nicaragua eliminated species of ancestral Hologymnetis previously living there. El gtnero Neotropical Hologymnetis (anteriormente Cineretis) es revisado. Se describen tres nuevas especies: H. kinichahau de Guatemala y MBxico, H. moronide Mtxico, y H. vulcanorum de El Salvador. Para cada especie se comentan 10s datos conocidos de su biologia y 10s registros precisos de localidad complementados con mapas y se incluye una clave para separar las siete especies hasta ahora conocidas. Se propone un anilisis cladistico que supone la filogtnia de este gtnero, discutiendo 10s caracteres y sus estados. Hologymnetis cinerea y H. undulata fueron encontradas las especies mis derivadas, mientras que H. vulcanorum, H. margaritis y H. kinichahau son menos derivados; H. moroni y H. argenteola son especies hermanas. Los estudios biogeogrificos indican que el gCnero es de origen sudamericano con una especie (H. undulata) ocumendo en el sur de la Cuenca del Amazonas. La dispersi6n a travis de Centroamtrica despues de la formacibn del puente panamefio es indicada. Los taxa descendientes se dispersaron y especieron en El Salvador a travts del sur y oeste de Mtxico y extremo sur de Arizona. L a subsecuente formaci6n de habitats desfavorables (primariamente bosques lluviosos en las tierras bajas) de Amazonia a Nicaragua elimin6 la especies ancestral de Hologymnetis anteriormente viva alli. 162 THE COLEOPTERISTS BULLETIN 46(2), 1992 The purpose of this revision is to provide a key for identification, description of adults, locality and temporal data, and the little biological information available for the species belonging to the Neotropical genus Hologymnetis. The genus Hologymnetis was previously composed of four species. We describe three new species here, two of which (the Guatemala and El Salvador species) were inadvertently discovered while studying possible variation in the male parameres of putative H. margaritis. Unlike related gymnetine genera such as Cotinis, Argyripa, and most Gymnetis, the form of the parameres is important in Hologymnetis because significant differences are present and consistently expressed. In spite of a bewildering array of colors and patterns in the Gymnetini, there are relatively few robust characters available to separate taxa from an otherwise widespread, nearly homogenous matrix of body form and sculpturing. Our cladistic study of these characters for Hologymnetis and their phylogenetic information is discussed in the section dealing with phylogeny. Lastly, the genus is geographically divided between Mesoamerica and central South America, and we postulate in the section on biogeography as to how this might have occurred. We believe the results of this study will now enable accurate and prompt identification of the species of Hologymnetis as well as add additional information to our efforts to understand the evolution and biogeography of scarabs. Traditional methods of alpha taxonomy were used to key and describe the species of Hologymnetis treated in this paper. Our database consisted of 1,692 specimens examined from the following institutions (acronyms are largely those used in Arnett and Samuelson 1986): (AMNH) American Museum of Natural History, New York (Lee Herman, Jr.); (BMNH) British Museum (Natural History), London (Michael Bacchus, Malcolm Kerley); (CASC) California Academy of Sciences, San Francisco (Norman Penny, David Kavanaugh); (CMNH) Carnegie Museum of Natural History, Pittsburgh (Robert Davidson, Ginter Ekis); (CNCI) Canadian National Collection, Ottawa (Ales Smetana, Jean McNamara); (DEIC) Deutsche Entomologisches Institut, Eberswald (H. Morge, H. Miiller); (FMNH) Field Museum of Natural History, Chicago (Henry Dybas); (IEMM) Instituto de Ecologia, Mexico City (now Xalapa) (Miguel Mor6n); (INPA) Instituto Nacional de Pesquisas da Amazonia (Norman Penny, Victor Py-Daniel); (KSUC) Kansas State University, Manhattan (Derrick Blocker); (LACM) Los Angeles County Museum, Los Angeles (Roy Snelling); (MCZC) Museum of Comparative Zoology, Cambridge (A1 Newton, Scott Shaw); (MHNG) Museum d'Histoire Naturelle, Geneva (Ivan Lobl); (MLUH) Martin Luther Universitat, Halle (Manfred Dorn); (MNHN) Museum National #Histoire Naturelle, Paris (Roger-Paul Dechambre); (MPEG) Museu Paraense Emilio Goeldi, Belem (William Overal); (MZSP) Museu de Zoologia, Slo Paulo (Cleide Costa); (QBUM) Museu Na~ional, Rio de Janeiro (Jost Carvalho); (SEMC) Snow Entomological Museum, Lawrence (George Byers); (UAIC) University of Arizona Insect Collection, Tucson (Carl Olson); (UCCC) Universidad de Concepcion, Concepcion (Tomas Cekalovic); (UCEC) University of Colorado, Boulder (U. Lanham); (UCRC) University of California, Riverside (Saul Frommer); (UNSM) University of Nebraska State Museum, Lincoln (Brett Ratcliffe); (USNM) United States National Museum, D.C. (Robert Gordon); 163 THE COLEOPTERISTS BULLETIN 46(2), 1992 and (ZMHU) Museum fur Naturkunde der Humboldt Universitat, Berlin (Manfred Uhlig, Fritz Hieke). Specimens were also examined from the following private collections: Dan Curoe, Palo Alto, CA; CuauhtCmoc Deloya, Xalapa, Mexico; John Glaser, Baltimore, MD; Alan Hardy, Sacramento, CA; Henry Howden, Ottawa, Canada; Paul Lago, University, MS; Antonio Martinez, Rosario, Argentina; Scott McCleve, Douglas, AZ,Miguel Morbn, Xalapa, Mexico; Brett Ratcliffe, Lincoln, NE; Carlos Seabra, Rio de Janeiro, Brazil; Don Thomas, Tuxtla Gutierrez, Mexico; and William Warner, Chandler, AZ. Field collecting was done by one or both of us in Brazil, Mexico, and the United States. We believe this to be an indispensible part of any revisionary study because it provides a greater understanding of the habitats, ecology, seasonality and actual abundance of the taxa under consideration. Specimens in this genus can usually be sexed using external characters. Males normally have flat or slightly concave sternites whereas females have rounded or convex sternites. Males usually have the central region of the sternites velutinous while females have this region shining and bare. The apical spurs on the posterior tibia in males are usually both acute, and in females both are usually rounded. Finally, the anterior tibia in males has a length/width ratio of 4: 1 whereas in females it is only 3: 1, hence relatively broader. It is important to note that the length measurements are from the anterior margin of the pronotum to the apex of the elytra. The head is not included in the measurement because it can be deflected at various angles upon preservation and, therefore, gives misleading indications as to overall length. Coloration and color terms are frequently difficult concepts with which to deal, especially in groups such as this one where many subtle shades of earth colors are present. Some of the color terms are absolutes and need no interpretation. For those terms that are open to misinterpretation based on one's own experiences, we have used Smithe (1 975) as a color standard in order to achieve objectivity. There are more complete color guides (e.g., Ridgeway 19 12), but they are not as widely available. The kind of light source is important when making color comparisons. Daylight, incandescent bulbs, and fluorescent tubes all give different color renditions, as does viewing with the unaided eye versus magnification. Except when stated otherwise, our use of color terms is made under magnification with incandescent scope lights striking the specimen at a 45" angle. The gazeteers published by the Central Intelligence Agency (1956a, b, 1965) were used to corroborate place names. The phylogeny analysis was conducted using cladistic methodology. The outgroup method of Watrous and Wheeler (198 1) was used to polarize characters into ancestral and derived states. PAUP (release 3), a phylogeny reconstruction program developed by David Swofford, was utilized in tree construction. Short of employing mitochondria1 DNA data, we believe that phylogeny analysis relying on phenotypic expression of characters is the best way to amve at relationship hypotheses. We are also acutely aware, however, that parallel and mosaic evolution, ecophenotypy, and small sample size all contribute to obfuscation of any hypothesis we might care to make using this methodology. Our biogeographical analysis is a straightforward use of vicariance and dispersal paradigms. It was also necessary to use a synthesis of data known for other organisms possessing a fossil record to aid in constructing dispersal patterns for our Scarabaeidae that do not have a fossil record. We have used a deductive approach to biogeographical reconstruction rather than narrative, inductive methods. "It is a normal practice in science to infer from what is 164 THE COLEOPTERISTS BULLETIN 46(2), 1992 better and more completely known in order to discover the structure and meaning of that which is less well or only partly known. The similarity of facts known on both sides of a controversy suggests that the best documented will be taken as a good model for the reconstruction of the structures still unknown on the other side" (Lavocat 1980:93). Plate tectonics, as exemplified by sea floor spreading and continental drift, is implicitly assumed in this study. Schiirhoff (1937) created the genus Cineretis for those species of Gymnetini lacking a strong, vertical, prosternal spine in combination with possessing three external teeth on the anterior tibia in both sexes. He included in this genus the species argenteola Bates, cinerea Gory and Percheron, and undulata Vigors. Unfortunately, he did not designate a type species for the genus (as required by Article 13b of the Code), and so the name is a nomen nudum. Martinez (1 949) recognized this and created a new genus, Hologymnetis, as a replacement name and designated Cetonia undulata Vigors as the type species. Hardy (1 975) correctly summarized these events. The Code's (Ride et al. 1985) definition of a nomen nudum is "a name that . . . if published after 1930, fails to conform to Article 13. A nomen nudum is not an available name and therefore the same name may be made available later for the same or a different concept; in such a case IT WOULD TAKE AUTHORSHIP AND DATE [emphasis mine] . . . from that act of establishment.. . ." Krikken (1984), in his treatment of the cetoniine genera of the world, was apparently unaware of Article 13b when he designated a type species (C. undulata) for Cineretis (of Schiirhoff) and declared Hologymnetis Martinez a junior synonym. According to Article 50g of the Code, Krikken became the author (though inadvertently) of Cineretis because "if a scientific name . . . is first published in the synonymy of an available name. . . its author is the person who published it as a synonym, even if some other originator is cited. . . ." Hardy (1 987) again correctly summarized this situation. Vigors (1825) described the first future Hologymnetis, Cetonia undulata, from Brazil. This species was then placed in the genus Gymnetis by Gory and Percheron (1 833). These authors also described Gymnetis consularis from Brazil, on a relatively immaculate specimen of G. undulata. This was correctly recognized as a junior synonym of G. undulata by Schenkling (1921) (based possibly on the advice of Moser). Gory and Percheron (1833) also described Gymnetis cinerea from Mexico. Blanchard (1 850) and Casey (1 9 15) then proceeded to describe an additional two and four species respectively that have since been placed in junior synonymy with G. cinerea (Blackwelder 1944). Examination of the types by one of us (Ratcliffe) confirms that they are all conspecific. Bates (1 889) described in the Biologia two additional species from Mexico, Gymnetis argenteola and G. margaritis. Casey (1 9 15) added, in his characteristic view of things, a new subspecies (laetula) from Arizona. Lastly, Blackwelder (1944) reduced G. margaritis to a subspecies of Cineretis argenteola. We do not adhere to this interpretation because of substantial differences between the taxa, and we restore margaritis to species status. Among the six syntypes (or possibly nine; see treatment for H. margaritis), there was one specimen not conspecific with the others; it is one of the paratypes of H. moroni described herein (while, at the same time, retaining its nomenclatorial paratype status for G. margaritis Bates). T H E COLEOPTERISTS BULLETIN 46(2), 1992 165 Hologymnetis Martinez Cineretis Schiirhoff 1937:56, 69 (nomen nudum). Hologymnetis Martinez 1949: 15. Type species: Cetonia undulata Vigors (by original designation). Cineretis Krikken 1984:60. Type species: Cetonia undulata Vigors. DESCRIPTION. Cetoniinae, Gymnetini. FORM: Rhomboidal, robust, sides tapering apically, dorsum weakly convex. Length from apex of pronotum to apex of elytra 11.522.0 mm; width across humeri 6.4-12.7 mm. COLOR: Varying on dorsum from opaque, velvety black to reddish brown to brown to cinnamon to olive gray or olive green to yellowish green to smoke gray to buff, with or without cretaceous pattern of indistinct fuscous cloudings. Ventrally, color varies from shining black to smoke gray with metallic copper reflections to metallic pale yellowish green or blue with or without copper reflections; sternites with or without cretaceous or contrasting color markings. HEAD: Shape subrectangular, longer than wide, weakly tumescent in center. Clypeus with apex slightly to distinctly emarginate and reflexed. Antenna with 10 segments, club with 3 segments not quite as long as stem. PRONOTUM: Widest at base, trapezoidal, attenuate apically, center-base produced into posteriorly projecting lobe. Lateral margin usually slightly sinuate before basolateral angle. ELYTRA: Widest at base, posthumeral emargination well developed; weakly elevated sutural costa and two short, discal costae terminating at apical umbone. PYGIDIUM: Surface transversely and concentrically strigate or coarsely and transversely vermiculate. VENTER: Vertical prosternal spine absent. Mesometasternal protrusion short to moderately long, ventral surface flat and in same plane as longitudinal axis of body. Males with sternites concave, each sternite with only a small, dark, shining patch or none. Females with sternites flat to slightly convex, sternites 2-5 in center with large, dark, shining patch. LEGS: Foretibia tridentate in both sexes, slender in males, wider in females. Metatibial spurs with apices acute in males, one or both rounded in females. DIAGNOSIS. The genus Hologymnetis may be distinguished from other New World Gymnetini by the following: head unarmed, prosternum lacking a long, vertical spine, foretibiae tridentate i n both sexes, and dorsum velutinous and largely unicolorous (except for the South American species that has distinctive cretaceous markings). RANGE. The combined ranges of the seven species extend from the southwestern United States to Honduras and from the southern half of Brazil to eastern Bolivia and southern Paraguay. 1. Venter largely shiny black in both sexes, sternites laterally often with cretaceous markings ............................................................................................................................... 2 1'. Venter brown to grayish brown or pale brassy green o r blue, weakly shining or weakly t o strongly metallic; females with central third of sternites bare, piceou 3 2. Dorsum unicolorous or black) (Fig. l) , rarely strongly punctate (Fig. 2). Mexico t o Guatemala (possibly Honduras) cinerea (G. & P.) 2'. Dorsum black o r piceous a lines; lines present longitudinally on pronotum and radiating from midline o n elytra (Fig. 5); occasionally reduced or nearly absent (Fig. 6). Southern Brazil, Paraguay, eastern Bolivia ............................ undulata (Vigors) 3. Venter brassy green (usually with coppery reflection) o r dull grayish olive or bluish gray (rarely blue), with o r without coppery reflections THE COLEOPTERISTS BULLETIN 46(2), 1992 3'. Venter grayish brown or yellowish brown, weakly shining 5 4. Venter a bright, shining brassy green, usually with distinc reflection. Protibia with teeth subequally spaced from one another. Parameres as in Figure 7. Arizona, northern Mexico ...... argenteola (Bates) 4'. Venter dull grayish olive or bluish gray (rarely blue), with or without coppery reflections. Protibia with median tooth distinctly closer to apical tooth th eres as in Figure 1 1. Westcentral Mexico moroni Ratcliffe & Deloya, n. sp. 5. Mesometastern ex usually sharply and transversely delineated from shaft (Fig. 16). Parameres with apical, median ridges tapering to a point (Fig. 13). El Salvador .. ..................... ................. ............. ....... ....... .................. vulcanorum 5'. Mesometasternal process with shining apex and shaft usually broadly joined by a br not acuminate 6 6. Parameres in pale, longitudinal median line. Southwestern Guatemala and Chiakinichahau Ratcliffe & Deloya, n. sp. 10). Pronotum with pale, longitudinal, median line. Southern Mexico margaritis (Bates) Hologymnetis argenteola (Bates) (Figs. 7 , 17) Gymnetis argenteola Bates 1889:354. Holotype male, labeled "Pinos Altos, Chihuahua, Mexico," "Sp. figured," "Buchan-Hepburn," "Type;" at BMNH (London), examined. Gymnetis argenteola laetula Casey 19 15:280. Type locality: "Nogales, Arizona." Holotype at USNM, examined. DESCRIPTION. Length 13.5-20.5 mm; width across humeri 7.6-1 1.1 mm. Color dorsally varying from entirely velvety dusky brown to clay to olive gray to olive green to yellowish green; vermiculate impressions and punctures shining black, piceous, or metallic pale green; ventrally metallic pale yellowish green with strong copper reflection, at least rim of punctures or strigae shining black. Occasional specimens with distinct cobalt blue suffusing pale green of venter. Other specimens lacking strong metallic lustre, instead weakly shining pale yellowish green or copper (depending on angle of light). HEAD: Surface moderately to moderately densely punctate; punctures small to moderately large, round to nearly crescent-shaped, with minute, pale setae in fresh specimens; declivous sides of clypeus rugulose. Clypeus feebly tumescent at center; apex strongly reflexed, usually strongly arcuate (best seen in anterior view). Interocular width 5.0-6.0 transverse eye diameters as seen from above. Antenna with basal segment at apex with dorsal fringe of 10 or fewer pale, slender setae. PRONOTUM: Surface usually moderately densely punctate; punctures small to large (largest in lateral one-third of pronotum), small punctures round, larger punctures crescent-shaped, setigerous in fresh specimens; setae short, pale; some specimens with entire field of punctures smaller throughout. Anterior margin lacking tubercle. Lateral border with complete marginal line, weakly emarginate before basolateral angle. Mesepimera moderately to densely punctate or rugopunctate; punctures moderate to large, round to vermiculate, setigerous; setae pale, often dense. ELYTRA: Surface moderately to densely punctate (denser in center of disc); punctures small to large, round to transverse to U-shaped, some setigerous; setae short, pale. Weakly elevated sutural costa and two discal costae terminating at apical umbone usually present in posterior half of elytra. PYGIDIUM: Surface coarsely, transversely vermiculate; impressions moderate to dense (occasionally confluent), C-shaped to elongate, setigerous; setae THE COLEOPTERISTS BULLETIN 46(2), 1992 167 Figs. 1-2. Habitus of H. cinerea. Fig. 1 (left) is typical. Fig. 2 (right) shows extreme in dorsal sculpturing. short, pale. Discal area flattened in both sexes. VENTER: Setae tawny. Meso-metasternal protrusion short, extending anteriorly to just beyond mesocoxae; anterior and ventral faces acute relative to one another, but angle rounded; ventral face flat, in same plane as longitudinal axis of body; anterior face with long, slender setae that curl around apex of protrusion. In ventral view, sides of mesometasternal protrusion diverging to broadly rounded apex. Last sternite at apex usually with fringe of slender, short, testaceous setae. Sternites completely yellowish green or with central area variably lacking green, instead shining black; punctures moderate to large, C-shaped to crescent-shaped, moderately dense laterally, becoming sparser medially. LEGS: Colored similarly to venter. Anterior tibia with 3 subequally spaced teeth. PARAMERES: Figure 7. DISTRIBUTION. Hologymnetis argenteola is found on the western slopes of the Sierra Madre Occidental in northwestern Mexico as well as in its northern extension, the Huachuca and Santa Rita mountains of extreme south-central Arizona in the United States. L o r n RECORDS (Fig. 17). 82 specimens examined. Specimens were seen from the following collections: AMNH, BMNH, CASC, CNCI, DEIC, LACM, MCZC, MFNH, MNHN, SEMC, USNM, ZMHU, Alan Hardy, Scott McCleve, Henry Howden, Miguel Morbn, Brett Ratcliffe, and William Warner. MEXICO (26): CHIHUAHUA (2): Moris, Pinos Altos; DURANGO (2): Canelas, Durango; SINALOA (2): NO data; SONORA (5): Agua Zarca, La Chiripa, 9 mi. NE Imuris, Nogales, Rio Mayo; No DATA (15). UNITED STATES (54): ARIZONA (48): COCHISE CO. (41): Carr Canyon, Chiricahua Mts., Coronado National Memorial, Hereford, Huachuca Mts. 2 mi. E. Nicksville, Palmerlee; PIMA CO. (1): 168 THE COLEOPTERISTS BULLETIN 46(2), 1992 Madera Canyon; SANTA CRUZ CO. (6): Nogales, Peiia Blanca Canyon (Oro Blanco Mts.), Santa Rita Mts. No DATA (9). TEMPORAL May DISTRIBUTION. (2), July (32), August (1 7). REMARKS. Hologymnetis argenteola is distinctive because of its unicolorous dorsum (similar to but usually greener and paler than H. cinerea) and venter metallic pale yellow green with strong copper reflections. The parameres are substantially different from those of H. cinereabecause the apex is more strongly toothed and the shaft of each paramere strongly curved (Fig. 7). Some slight variation was observed in the slenderness in the middle of each paramere. The dorsal coloring and degree of punctation of this scarab varies considerably with no apparent correlation to geography or altitude. So, too, does the ventral coloration. Metallic pale yellowish green with strong coppery reflections is most common. Some specimens have metallic bluish green reflections beneath. Bates' holotype is similar to this, as is the holotype of Casey's laetula. The presence of a distinct canna ventrally and laterally near the base of each paramere (Fig. 7), in combination with the form of the apices of the parameres and geographic origin, will distinguish this species. The ventral color of these two species is also different but is not as reliable a character for separation. Biological information is sparse for H. argenteola, but presumably the adults are flower and sap feeders and the larvae develop in organic debris or compostlike situations. Label data indicate that some specimens were collected from gumming sites on Baccharis sararthroides Gray, from banana traps and from "desert willow" at elevations ranging from 1,212-2,273 m. Adults have occasionally been found competing with species of Euphoria and Cotinis at gumming sites on Baccharis sp. (William Warner, pers. comm. 1989). This species is apparently not abundant. Hologymnetis cinerea (Gory and Percheron) (Figs. 1-2, 8, 17) Gymnetis cinerea Gory and Percheron 1833:73, 37 1. Lectotype male, labeled "Mexico" here designated; at MHNG (Geneva) with Ratcliffe's lectotype label. Also two male paralectotypes labeled "Mexico" at Geneva and one female paralectotype labeled "Mexico, Gory," "Coll. Helfer," "Typus," "Exemplaire typique du cabinet de M.H. Gory," and "Field Mus. Coll., F. Psota Coll. (ex. Aug. Ondrej Coll.)" at FMNH (Chicago). Gymnetis punctata Blanchard 1850:36. Type locality: "Mexique." Holotype at MNHN (Paris), examined. Gymnetis uniformis Blanchard 1850:36. Type locality: "Mexique." Holotype at MNHN (Paris), examined. Gymnetis cuneata Casey 19 15:282. Type locality: "Durango City, Mexico." Holotype and allotype at USNM (Washington, D.C.), examined. Gymnetis lobiculata Casey 19 15:282. Type locality: "Mexico, probably near Jalapa." Holotype at USNM (Washington, D.C.), examined. Gymnetis simulans Casey 1915:283. Type locality: "Mexico (Guerrero)." Holotype and allotype at USNM (Washington, D.C.), examined. Gymnetis aequalis Casey 19 15:284. Type locality: "Cuernavaca, Morelos, Mexico." Holotype at USNM (Washington, D.C.), examined. DESCRIPTION. Length 14.7-21.7 mm; width across humeri 8.5-12.7 mm. Color dorsally varying widely from entirely velvety or opaque black to dark gray to dark plum to light or dark cinnamon to grayish or brownish olive to light or dark smoke gray to buff; black where punctate or velvety surface absent (head occasionally entirely black); venTHE COLEOPTERISTS BULLETIN 46(2), 1992 trally shining, black. HEAD: Surface usually sparsely to moderately punctate either side of raised midline; punctures small to moderate in size, setigerous in fresh specimens; where velutinous covering absent, surface densely rubopunctate; declivous sides of clypeus rugulose, clypeal apex rugulose to finely punctate; setae tawny, short. Clypeus with apex reflexed, distinctly arcuate (curve bowed posteriorly) to subtruncate. Interocular width equalling 5.0-6.0 transverse eye diameters. Antenna with basal segment at apex with dorsal fringe of 10 or fewer pale or dark colored, slender setae. PRONOTUM: Surface usually completely velutinous, sculpturing obscured; otherwise, varying from sparsely punctate in anterior half with punctures small and round to moderately densely punctate all over with punctures small, round mixed with moderate to very large round, crescentshaped or U-shaped punctures. Anterior margin with small tubercle. Lateral margin completely beaded, not or only weakly emarginate before basolateral angle. Mesepimera largely velutinous, usually with a few small to moderate punctures in anterior half. ELYTRA: Surface entirely velutinous, punctation varying from absent to sparsely to moderately punctate; punctures varying from small and round (in which case punctures usually in partial lines representing striae and random in intervals between striae) to moderate and large mixed; larger punctures crescent shaped (in which case, punctation seemingly random). Weakly elevated sutural costa and 2 discal costae terminating at apical umbone usually present in posterior half of elytron. PYGIDIUM: Surface usually transversely, concentrically strigulose around a point at center apex; otherwise (in specimens with coarse pronotal and elytral sculpturing) surface transversely strigate in basal half, apical half with sparse, transverse, moderate to large punctures; strigae and/or punctures setigerous; setae short, pale, dense at base, becoming sparser apically (some specimens seen with dense setae over entire surface). In some specimens surface velutinous, obscuring sculpturing. VENTER: Setae testaceous to mostly black. Mesometasternal protrusion moderately long, extending well past mesocoxae, but not quite reaching forecoxae; anterior and ventral faces form sharply acute angle. Ventral surface flat, in same plane as longitudinal axis of body; anterior face with short, stout setae. In ventral view sides subparallel to slightly divergent to broadly rounded apex. Sternites 1-5 occasionally with small area of velvety color (as on dorsum) on lateral margin and apical edge of each segment (usually seen only in specimens with pygidium similarly colored). LEGS: Markings absent, color piceous to black, shining. Anterior tibia with 3 subequally spaced teeth, basal tooth often reduced. PARAMERES: Figure 8. DISTRIBUTION. Except for the Baja peninsula and extreme north-central Mexico, H. cinerea is found throughout Mexico west of the isthmus of Tehuantepec. There are several records east of the isthmus in Chiapas but not in the Yucatan peninsula. There are few records for Guatemala and questionable records for El Salvador, Honduras, Arizona, and Texas. LOCALITYRECORDS (Fig. 17). 1,238 specimens examined. Specimens were seen from the following collections: AMNH, BMNH, CASC, CMNH, CNCI, DEIC, FMNH, IEMM, KSUC, LACM, MCZC, MHNG, MLUH, MNHN, MCSP, SEMC, VCEC, VCRC, UNSM, USNM, ZMHU, John Glaser, Alan Hardy, Henry Howden, Antonio Martinez, Miguel Morbn, Cuauhthmoc Deloya and Brett Ratcliffe. EL SALVADOR (1): Cafetalera. (1). GUATEMALA: (2 1) BAJA VERAPAZ (2): Candiza, (16): San Geronimo, No data; CHIQUIMULA Chiquimula; SANTA ROSA (3): Barcena. HONDURAS (3) No DATA (3). MEXICO (1,212): AGUASCALIENTES (3): Santa Rosa (5 ) : Aguascalientes; CHIAPAS (Lagunas de Montebello), Sumidero Cnyn. Nat. Pk., No data; CHIHUAHUA (3): San Isidro, Temosachic; COLIMA (7 1): Colima, Tonila, Volcan de Colima, No data; DF (22): Mexico City, Temascaltepec, No data; DURANGO (35): Durango, 5 mi. W Durango, 18 mi. W Durango, Nombre de Dios, Ventanas, Villa Lerdo, No data; GUANAJUATO (9): Acambaro, 11 mi. SW Acambaro, No data; GUERRERO (41): Acahuizotla, Acapulco, Amuco de la Reforma, Chilpancingo, 3 mi. N Chilpancingo, 10 km E Chilpancingo, Cuacoyula, Taxco, 3 mi. N 170 THE COLEOPTERISTS BULLETIN 46(2), 1992 Taxco, Teloloapan, Tierra Colorada, La Venta, 7 km E Xochipala, No data; HIDALGO(4): Barranca Venados, Melchor Ocampo, No data; JALISCO (229): Ajijic, Chapala, Chamela Biol. Sta., 12 mi. S Encarnaci6n de Diaz, Guadalajara, 8 and 10 and 23 mi. S Guadalajara, 28 and 53 mi. E Guadalajara, 7 and 16 and 19 and 25 mi. W Guadalajara, Huejotitan, Jacotopec, 6 mi. NE Jolostitlin, Lagos de Moreno, 15 mi. NE Lagos de Moreno, 13 mi. SE Lagos de Moreno, 22 mi. NW La Piedad, 20 mi. N La Quemada, Ocotlin, 8 mi. W San Juan de 10s Lagos, 8 mi. SW San Juan de 10s Lagos, 9 mi. W Tepatitlan (El Refugio), El Tuito, 10 mi. E Union de Tula, Zapopan; MEXICO (5) : Canalejas, Ixtapan de la Sal, Tejupilco; MICHOACAN (13): Acahuato, Cotija, Huetamo, Jungapco, 10 mi. N Morelia, 4 mi. W Morelia, Palo Alto, Tacambaro; MORELOS (127): Caiiada de Lobo, Cuautla, 10 mi. W Cuautla, Cuernavaca, Jojutla, Morelos, Puente de Ixtla, Tepoztlan, Tequesquitengo, Tlaltizapan, Tlaquiltenango, Yautepec, Xochicalco, Krn 6 on Hwy 160 from Jonacatepec to Atotonilco; NAYARIT (19): Compostela, 18 mi. W Compostela, Jeshs Maria, 40 km SE Peiiitas, 5 mi. S Rio Santiago ferry, Tepic, 24 mi. SE Tepic, Tuxpan; N u ~ v o LEON (8): 15 mi. SW China, Monterrey, 5 mi. S and 20 mi. S and 50 mi. S of Monterrey, Villa Santiago; OAXACA (6 1): La Asunci6n (1 2 mi. NW Oaxaca), Etla, Juquila Mixes, Mitla, Oaxaca, 10 mi. S Oaxaca, Tamaulapan, 56 mi. NW Tehuantepec, Teotitlan Tlacolula, No data; PUEBLA (64): Acatlan, Atlixco, Guadalupe, Izhcar de Matamoros, Tehuacan, 6 mi. NE Tehuacan; QUER~TARO (2): NO data; SAN LUIS POTOSI (4): El Naranjo, Ciudad Valles, San Luis Potosi; SINALOA (31): 5 mi. NW Choix, 13 mi. E La Concordia, 17 km E La Concordia, Mazatlan, 5 mi. N Mazatlan, Presidio, Venodio, Villa Union; SONORA (48): Alamos, 7 mi. and 10 mi. S Alamos, Arispe, Estrella Alamo, Guirocoba, 15 mi. E Navajoa, San Bernadino (Rio Mayo), Ventanas, No data; VERACRUZ (64): Acatlan, Cordoba, Cotaxtla, Jalapa, Misantla, Orizaba, Plan del Rio, Sierra Mixteca, Tierra Blanca, Tuxpan, Veracruz, No data; ZACATECAS (2): Monte Escobido, Tlaltenango de Sanchez Rom6n; No DATA (373). UNITED STATES (2): ARIZONA (1): No data; TEXAS (1): NO data. TEMPORAL January (I), March DISTRIBUTION. (I), April (1), May (I), June (22), July (85), August (103), September (123), October (68), November (1 8), December (4). REMARKS. Hologymnetis cinerea, in spite of its wide variety of color forms, is easily distinguished from its congeners by the presence of both a unicolorous dorsum and a shiny black venter. The parameres of the male genitalia were seen to vary in the degree to which the subapical flanges or "teeth" were produced. In most specimens, they are produced (Fig. 8) while in others they are not raised appreciably from the surface plane of the paramere. This difference did not correlate with any altitudinal or geographic parameters. Similarly, some specimens are coarsely punctate on the pronotum and elytra (Fig. 2). As Bates (1 889) noted, punctation is present in certain specimens of all the color forms and is seemingly of little taxonomic importance. We concur with this, having noted no correlation between the degree of punctation and locality or altitude. This species is often locally abundant feeding on flowers. It has been taken feeding on Acacia angustissima (Mill.) and Baccharis glutinosa Pers. in Nayarit and Jalisco states. Adults have also been found in the detritus piles of leafcutter ants (Atta sp.) (Deloya 1988) in Hidalgo, Morelos, and Nayarit states and have been taken in fruit-baited traps in Guerrero. Hologymnetis cinerea has a broad tolerance of different habitats inasmuch as it has been found from the lowland forests (150 m elev.) of Veracruz to the open pine highlands (2,000 m elev.) of Michoacan. THE COLEOPTERISTS BULLETIN 46(2), 1992 Hologymnetis kinichahau Ratcliffe and Deloya, n. sp. (Figs. 3, 9, 15, 17) TYPE MATERIAL. Holotype labeled, "GUAT., Finca San Rafael Olimpo, Cuyotenango, Such., X101965, J.M. Campbell, 1700." Allotype labeled, "MEXICO: Chiapas, San Cristobal de las Casas, VII-1988, A. Mor6n col." Five paratypes with the following data: "MEXICO: Chiapas, Municipio Motozintla, Ridge between Cerro Boqueron & Niquivil, 2438-2743 M, 15-XII1976, D.E. & J.A. Breedlove, Cal. Acad. Sci. Coll." (1); "Rincon Chamula, Chis., MEX. 7-VII-1969, L.A. Kelton" (1); "Chiapas, MEXICO, San Cristobal, 22-23 Jun. 1989, P.K. Lago" (1); "MEXICO, Chiapas, San Quintin, 27-VII1978, P. Hubbell" (1); "MEXICO, Chiapas, Rancho Santa Rosa, 17-XI1975, P. Hubbell" (1). Holotype at CNCI. Allotype in the collection ofMiguel Mor6n. Paratypes at CASC, UAIC, and in the Paul Lago and Brett Ratcliffe collections. HOLOTYPE. Male: Length 17.2 mm; width across humeri 10.6 mm. Color dorsally an opaque brown to olive brown, punctures and sculpturing piceous to black; venter weakly shining, light brown to darker smoke gray with faint coppery reflection; sculpturing black. HEAD: Surface moderately densely punctate; punctures small on frons, becoming large on clypeus, round to oblong; setae not present; declivous sides of clypeus rugulose. Clypeus slightly convex on disc; apex broadly reflexed, distinctly emarginate at center, apices either side of emargination broadly rounded. Interocular width equaling 5.0 transverse eye diameters as seen from above. Antenna with basal segment at apex with fringe of 12 testaceous, slender setae on dorsal edge. PRONOTUM: Surface with sparse, minute punctures on center of disc, becoming moderately dense on lateral thirds of pronotum where punctures large, round, deep. Anterior margin at center not tumid. Lateral margin with bead broken up by sculpturing; side margin just anterior of basolateral angle distinctly emarginate in dorsal view. Mesepimera moderately punctate; punctures large, round, deep, setigerous; setae short, testaceous, most apparently abraded away. ELYTRA: Surface sparsely punctate on disc, becoming denser behind apical umbones; punctures large, mostly round on disc to mostly kidney-shaped apically. Sutural costae weakly elevated. PYGIDIUM: Surface coarsely, densely punctate/vermiculate; impressions kidneyshaped to transverse to elongate, deep, setigerous; setae short, testaceous. VENTER: Setae testaceous. Mesometasternal protrusion short, extending anteriorly just beyond mesocoxae; anterior and ventral surfaces weakly acute relative to one another, angle rounded; ventral surface flat, in same plane as longitudinal axis of body; anterior surface with long, slender setae that curl around apex of protrusion. In ventral view, sides of mesometasternal protrusion diverging to broadly rounded apex. Sternites with large punctures in lateral thirds; punctures moderately dense, round to mostly kidney-shaped, setigerous; setae short, testaceous. LEGS: Colored similarly to venter. Anterior tibia with 3 subequally spaced teeth. PARAMERES: Figure 9. ALLOTYPE. Female: Length 17.7 mm; width 10.4 mm across humeri. As holotype except in following respects: Color light brown; elytra with broad, transverse, diffuse lighter yellowish brown band just behind middle (Fig. 3); venter light brown with weak metallic reflection. ELYTRA: Punctures very large, about twice size of those in holotype, U-shaped to C-shaped with opening facing posteriorly. VARIATION. Males (2 paratypes): Length 19.8-21.9 mm; width across humeri 12.012.9 mm. Color and pattern as allotype. Other than the "freshness" of the specimens (reflected in distinct color and pattern), there is little variation from the holotype. The three teeth on the anterior tibiae in both specimens are nearly obsolete. Females (3 paratypes): Length 6.1-9.0 mm; width across humeri 9.9-1 1.2 mm. Color as allotype (1 specimen) to darker brown (2 specimens). HEAD: Punctation a little denser in one specimen. ELMRA: Punctures large and round in one specimen as opposed to C-shaped. DISTRIBUTION. Hologymnetis kinichahau is known only from the Pacific slope of the Sierra Madre in Guatemala and Chiapas state in Mexico (Fig. 17). 172 THE COLEOPTERISTS BULLETIN 46(2), 1992 Fig. 3. Habitus of H. kinichahau. 173 THE COLEOPTERISTS BULLETIN 46(2), 1992 REMARKS. Hologymnetis kinichahau is similar in external appearances to H. margaritis. Whereas there is a lighter, longitudinal line on the pronotum of H. margaritis, there is no such line in H. kinichahau. If anything, this part of the pronotum is a little darker with an irregular, fuscous clouding. The parameres of the male genitalia are distinctive because they are so short and stout. ETYMOLOGY. Being a diurnal species that is most active in bright sunlight, we name this species after the sun god, Kinich Ahau, of the ancient Maya indians of Guatemala. Hologymnetis margaritis (Bates) (Figs. 4, 10, 14, 18) Gymnetis margaritis Bates 1889:354. Lectotype male, labeled "Tehuantepec," "B.C.A., Col. I1 (2) ,Gymnetis," "Type, sp. figured," "Mexico, Salle coll.," and "B.C.A. figured," here designated; at BMNH (London) with Ratcliffe's lectotype label. Female lectoallotype, labeled "Acapulco, Guerrero," "B.C.A., Col. I1 (2), Gymnetis," and "Hoge" here designated; at BMNH with Ratcliffe's lectoallotype label. Also, one male paralectotype labeled as lectoallotype, one male paralectotype labeled "Guatemala," "margaritis var.," "Salle coll.," and one unsexed paralectotype labeled "Oaxaca, Mexico, Hoge," "B.C.A., Col. I1 (2) , Gymnetis" here designated; all at BMNH with Ratcliffe's paralectotype labels. One additional male paralectotype, labeled "Oaxaca, Mexico, Hoge," "B.C.A., Col. I1 (2), Gymnetis margaritis Bates," and "Cotype CNCNo. 8676, G. margaritis Bates" at CNCI (Ottawa) with Ratcliffe's paralectotype label. One specimen labeled "Acapulco" and two labeled "Etla" are at MNHN and may be part of the original type series because their label data match those in the original description. Bates indicated there was only one specimen from Etla, however, and did not mention numbers for Acapulco. Due to these uncertainties, lectotype designations were not made for the specimens at Paris. DESCRIPTION. Length 11.6-17.2 mm; width across humeri 6.4-10.2 mm. Color dorsally varying from opaque light to dark brown; pronotum usually with light brown, narrow, longitudinal line in center; elytra usually with broad, transverse, d~fuse light brown 'band' just behind middle (Fig. 4) and usually with weakly raised costae (terminating in apical umbone) darker than background color (not noticeable in dark brown specimens); venter weakly shining, darker smoke gray, occasionally with faint metallic copper reflection; punctures and other sculpturing piceous to black. HEAD: Surface moderately densely to occasionally densely punctate; punctures small to large (frequently larger on frons than on clypeus), round to kidney-shaped, setigerous; setae minute, straw colored; declivous sides of clypeus rugulose. Clypeus feebly tumescent at center; apex strongly reflexed, distinctly emarginate at center, apices usually distinctly angulate either side of center. Interocular width 4.0-5.0 transverse eye diameters as seen from above. Antenna with basal segment at apex with 10 or fewer pale, slender setae on dorsal edge. PRONOTUM: Surface moderately (most common) to densely punctate; punctures small to larger (larger and denser laterally), round to kidney-shaped, setigerous in fresh specimens; setae minute, pale. Anterior margin at center not tuberculate or noticeably timid. Bead of lateral margin varying from complete to broken or missing in basal half; margin just anterior of basolateral angle varying from entire to distinctly emarginate. Mesepimera moderately to densely punctate; punctures small to mostly large, round to crescent-shaped to vermiculate, setigerous; setae pale, often dense, long. ELYTRA: Surface moderately to densely punctate (much less so basomedially); punctures varying from small to mostly large, round to transverse to kidney-shaped to U-shaped to vermiculate near apex. Weakly elevated sutural costa and 2 discal costae terminating at apical umbone present in posTHE COLEOPTERISTS BULLETIN 46(2), 1992 Fig. 4. Habitus of H. margaritis. THE COLEOPTERISTS BULLETIN 46(2), 1992 175 terior half of elytra. PYGIDIUM: Surface coarsely, transversely vermiculate (rarely with large, transverse punctures only); impressions moderate to dense (occasionally confluent), C-shaped to elongate, setigerous; setae short, pale. VENTER: Setae testaceous. Mesometasternal protrusion short, extending anteriorly beyond mesocoxae but not reaching procoxae; anterior and ventral surfaces weakly acute relative to one another, but angle rounded; ventral surface flat, in same plane as longitudinal axis of body; anterior surface with long, slender setae that curl around apex of protrusion. In ventral view, sides of mesometasternal protrusion slightly diverging to rounded apex. Apex of protrusion piceous, shining, broadly connected by similar colorations to shaft of protrusion (Fig. 14). Last sternite at apex with fringe of slender, short, testaceous setae. Sternites completely smoke gray or with small piceous spot basomedially (males) or with central area variably shining black (females); punctures moderate to large (larger laterally), kidney-shaped to C-shaped. LEGS: Colored similarly to venter. Anterior tibia with 3 subequally spaced teeth. PARAMERES: Figure 10. DISTRIBUTION. Hologymnetis margaritis is found in the Sierra Madre del Sur in southwestern Mexico and in the Sierra Madre of southern Chiapas. There is a country record only for El Salvador. L O C A L I ~ (Fig. 18). 53 specimens examined. Specimens were seen RECORDS from the following collections: AMNH, BMNH, CNCI, MNHN, SEMC, USNM, ZMHU, Henry Howden, CuauhtCmoc Deloya, Miguel Morbn, Don Thomas, and Brett Ratcliffe. GUATEMALA (1): No DATA (1). MEXICO (5 1): CHIAPAS (4): Ciudad CuauhtCmoc, El Aguacerro, Tapachula; GUERRERO (9): Acahuizotla, Acapulco, Amula, No data; JALISCO (2): Chamela Biol. Sta., 10 km N W El Tuito; MICHOACAN (4): 14 mi. E San Blas, Tepic; (1): Uruapan; NAYARIT OAXACA(21): Ayoquesco, Etla, Hiway 190 at Chiapas border 19 mi. S Matias Romero, Monte Alban, Oaxaca, Pochutla, San Mateo Macuilzochitl Tehuantepec, 56 mi. NW Tehuantepec; No DATA (9). TEMPORAL May DISTRIBUTION. (1), June (1 5), July (8), October (3), November (1). REMARKS. Hologyrnnetis margaritis, although variable in its dorsal markings and color, is nearly always characterized by having an irregular, lighter band across the elytra just past the middle (best seen by the unaided eye). Only three of the 45 specimens studied lacked this band. Frequently, this band is delimited posteriorly by the darker, raised costae terminating at the apical umbone. Normally, the pronotum has a lighter, longitudinal, narrow band at its center that is absent in the other species of the genus (best seen with magnification). The ventral coloring of smoke gray or grayish brown is also distinctive. Little variation was observed in the form of the parameres, which are also unique. They resemble most closely those of H. argenteola but lack the apicolateral tooth. Life history information for this species is lacking. Label data indicate that specimens have been taken at elevations ranging from 150 m at Tapachula, Chiapas to 1,936 m at Monte Alban, Oaxaca. Hologymnetis moroni Ratcliffe and Deloya, n. sp.