MSU LIBRARIES J-I-I1llll. ‘v _....’.w__ .._.__. RETURN I NG F'?/"«.T‘ERU3\[_S: Place in book drop to remove this checkout from your record. [gggg wi11 be charged if book is returned after the date stamped be1ow. monomcs, PHYLOGENY AND sys'rmn'ncs or rm: NORTH AMERICAN spac- or PEDILUS mscmm (cowor'rmm: rYnocanomAB) by Daniel K. Young A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Entomology 1981 / - P" O f 2' // 1/ / r f I ‘ ~ ‘ A y 31011011103, PHYLOGENY AND sxs'rRMA'ncs or ran NORTH AMERICAN spac- or PEDILUS FISCHER (COLBOPTERA: PYROCRROIDAB) by Daniel K. Young The thirty North American species of the genus Pedilus are reviewed. Bionomics as well as suprageneric and intrageneric relationships are discussed in some detail and a number of hypotheses relative to the natural history and phylogeny of the gems are presented. Larvae are associated with decaying vegetative materials at or near the soil surface while adults are typically found on flowers. Thus far, males of twenty species have been shown to orient positively toward cantharidin, the chemical defense mechanism of meloid and some oedemerid beetles. Evidence from an analysis of adult and available larval stages indicates that Pedilus is more closely related to the Pyrochroidae than to either the Eurygeniinae or the Anthicidae (sensu stricto); the Pedilidae and Anthicidae (sensu lato) are not tenable. Taxonomic history, methodology and general anatomy are discussed and diagnostic keys to adult males and females are presented. The North American faunal element of Pedilus consists of six species groups with two species unplaced and one fossil species of uncertain disposition. All taxa are described or redescribed along with the following: synonymies, geographic distribution, materials examined, diagnoses, and general remarks; pertinent diagnostic features are illustrated. One new synonym is added at the generic level along with thirteen at the species level; four new species are described: fenderi, granti, joanae, and johnsonorum. ii ACKNOWLEDGMEN'IS Most of the 11,585 specimens of North American Pedilus which formed the data base for this study were borrowed from the following curators and private collectors: (AMCC) - A.J. Mayor, Riverside, CA [Private]; (AMNH) - L.H. Herman, American Museum of Natural History; (BMNB) - C.M.F von Hayek and R.J.W. Aldridge, British Museum of Natural History, ,London, England; (BPBM) - G.A. Samuelson, Bernice P. Bishop Museum; (CASC) - D.H. Kavanaugh, California Academy of Sciences; (001:0) - 0. Chantal, Quebec, 9.0. [Private]; (CCNY) - D.C. Miller, City College of New York; (CDAB) - F.G. Andrews, California Department of Food and Agriculture; (CISC) - J.A. Chemsak, University of California, Berkeley; (CNCD - E.C. Becker, Canadian National Collection of Insects; (CUCC) - J.C. Morse, Clemson University; (CUIC) - L.L. Pechuman, Cornell University; (DBUM) - F. Harper, University of Montreal; (DBFW) - P.J. Clausen, University of Minnesota; (DEUN) - H.L. Gunderson, University of Nebraska; (DSCC) - D.S. Chandler, Durham, NH [Private]; (D821) - R. Baranowski, Department of Systematics, Zoological Institute, Lund, Sweden; 111 (DYCC) - collection of the author; (DZBC) - N.L. Anderson and S. Rose, Montana State University; (BJKC) - E.J. Kiteley, Quebec, P.Q. [Private]; (BIDS) - W.J. Hanson, Utah State University; (EEOC) - E.G. Riley, Columbia, MO [Private]; (PINE) - the late H. Dybas and L.E. Watrous, Field Museum of Natural History; (PSCA) - R.R. Woodruff, Florida Department of Agriculture; (GHNC) - G.H. Nelson, Pomona, CA [Privatd; (INKS) - D.W. Webb, Illinois Natural History Survey; (JBJC) - J.B. Johnson, Moscow, ID [Privatd; (JKLC) - J.K. Liebherr, Berkeley, CA [Private]; (JSCC) - the late J. Schuh, Klamath Falls, OR [Private] (collection transferred to American Museum of Natural History); (KSUC) - H.D. Blocker, Kansas State University; (LACK) - C.L. Hague, Los Angeles County Museum; (LSUC) - J.B. Chapin, Louisiana State University; (MCZC) - F.M. Carpenter, Division of Fossil Insects, Museum of Comparative Zoology; (MCZC) - J.F. Lawrence and A.F. Newton, Museum of Comparative Zoology; (INKS) - R.R. Hooper, Saskatchewan Museum of Natural History; (MSUC) - R.L. Fischer, Michigan State University; (NAUF) - C.D. Johnson, Northern Arizona University; (NCSU) - D.L. Stephan, North Carolina State University; (NDSU) - E.U. Balsbaugh, North Dakota State University; (NHMB) - W. Wittmer, Naturhistorisches Museum, Basel, Switzerland; (NMDC) - N.M. Downie, Lafayette, IN [Private]; 1v (NMPC) - S. Bily, National Museum, Prague, Czechoslovakia; (ODAC) - R.L. Westcott, Oregon Department of Agriculture; (OSUC) - C.A. Triplehorn, Ohio State University; (0800) - B.B. Frost, Oregon State University; (0800) - S. Rohwer, Burke Museum, University of Washington (collection transferred to Oregon State University); (PADA) - T.J. Henry and A.G. Wheeler, Jr., Pennsylvania Department of Agriculture; . (PMNH) - C.L. Remington, Peabody Museum, Yale University; (PSUC) - K.C. Kim, Pennsylvania State University; (PURC) - A. Provonsha, Purdue University; (RAWC) - R.A. Wharton, San Francisco, CA [Private]; (Rnwc) - R.D. Ward, Washington, D.C. [Private]; (ROMC) - G.E. Wiggins, Royal Ontario Museum; (SHIN) - S.E. Miller, Santa Barbara Museum of Natural History; (SBMC) - P.D. Ashlock and G.W. Byers, Snow Entomological Museum, University of Kansas; (SMCC) - S. McCleve, Douglas, AZ [Private]; (TAMU) - H.R. Burke, Texas A&M University; (TWTC) - T.W. Taylor, Ft. Davis, TX [Private]; (UADB) - E.P. Rouse, University of Arkansas; ,(UASM) - D. Shpeley and G.E. Ball, University of Alberta; (UBCZ) - J.A. van Reenen, University of British Columbia; (UCDC) - R.O. Schuster, University of California, Davis; (UCEC) - U.N. Lanham, University of Colorado; (UCRC) - S.I. Frommer, University of California, Riverside; (UGCA) - R.H. Turnbow and C.L. Smith, University of Georgia; (ULIC) - C.V. Covell, Jr., University of Louisville; (UMMZ) - LJ. Cantrall, University of Michigan; (DIRK) - W.R. Enns and EG. Riley, University of Missouri; (USNM) - T.J. Spilman, United States National Museum of Natural History; (UVCC) - R.T. Bell, University of Vermont; (VPIC) - M. Kosztarab, Virginia Polytechnic Institute; (WHCC) - W.R. Clark, Boise, ID [Private]; (wsCC) - w.s. Craig, Columbia, MO [Private]; (WSUC) - W.L. Turner, Washington State University Special thanks go to the following individuals who offered their assistance with cantharidin baiting, donated their personal material to the author or provided helpful comments throughout the course of the study: A.D. Allen, D.S. Chandler, C. Chantal, K.M. Fender, R.L. Fischer, D. Flynn, R.R. Hooper, J. Ingerson-Mahar, J.B. and J.T. Johnson, P.J. Johnson, H.J. Lee, Jr., J.K. Liebherr, S. McCleve, J.B. McPherson, S.E. Miller, M. Oemke, F.W. Ravlin, L. Russell, M.W. Sanderson, G.A. Shook, R.D. Ward, and 8.6. Wellso. John F. Lawrence kindly allowed the use of an illustration (Figure 133) prepared by Susan Poulakis; the rough version of the text and many of the tables were typed by Mrs. Lucille Wells. I am grateful to the members of my dissertation and program guidance committee: Drs. Ring T. Carde', Roland L. Fischer, Frederick W. Stehr, Charles A. Triplehorn, and Stanley G. Wellso. Their freely given time and energies, and interest in my professional development have been greatly appreciated. vi At numerous points throughout the course of my undergraduate and graduate programs, I have received much needed encouragement and love from friends and family members. To these persons, particularly my mother and father, I am most grateful. Finally, I wish to express my thanks to my dear wife, Joan. Her love and understanding have contributed so very much to the fruition of this project. vii TABLE OF CONTENTS LISTOFTABLES ............................... ..... . ...... .. x1 LIST OF ILLUSTRATIONS........ ..... .. ......... . ....... . ..... xiii INTRODUCTION.............................................. 1 BIONOMICS OF NORTH AMERICAN SPECIES OF Pedilus ........ 3 HISTORICALREVIEWFOR NORTH AMERICA.................... 10' METHOWLOGYOCOO00.00.000.000......IOOOOOOOOOO...00......O 15 CRITERIA FOR SPECIES RECOGNITION AND FOR INFRASPECIFIC AND SUPRASPECWIC GROUPINGS ......OOOOOCOOOCOOOOOOOO 30 GENERIC AND SUPRAGENERIC PHYLOGENETIC CONSIDERATIONS 32 INTRAGENERIC PHYLOGENETIC CONSIDERATIONS . . . . . . . . . . . . . 46 HISTORICAL BIOGEOGRAPHY ............................ ..... 74 CLASSIFICATION OF NORTH AMERICAN SPECIES . . . . . . . . . . ..... 79 CHECKLIST OF NORTH AMERICAN Pedilus SPECIES GROUPS ANDSPECIES ......OOOOOOOOOOOO......OOOO...0.0.0.000... 89 KEYTOADULTMALES.................... ............. . ..... 91 KEYTOADULTFEMALES(REGION1)... 99 KEY TO ADULT FEMALES (REGION 2).. ..... ...... . ......... 101 KEY TO ADULT FEMALES (REGION 3) ........... . ........ ... 102 THEFLABELLATUSGROUPO..........OOOOOOOOOOOOOOOOOO...... 104 PedlmflaMllam(Hom)0000000000000000000.000 ........... 105 PedillerViCdlisPal-100000000000000000000000 ..... ... ..... 110 viii TABLE OF CONTENTS, continued THEPUNCTULATUS GROUPOOOOOOO......OOOOOOOOOOOO0.0.000... Pedilusabnormis (Horn) Pedilusdentatus Abdullah-------------------- .............. .Pedilusmsticus Abdullah--- Pedilusfenderi,NEwspECIEs..... ..... ..... Pedilusflexiventris Fall PediluscavatusFan.............. ..... ....... Pedilusiohnsonorwn,NEw SPECIEs......................... Pedilus inconspicuus (Horn) . .. ........ . .................... PedIIUSVittatus (Horn) ........................... ....... .. Pedilusoregonus Fall Pedilusbardi (Horn) Pediluspunctulatus LeConte THE LEWISI GROUP . ......................................... Pedilus picipennis Fall ..................................... Pedflus elegans (Hentz) .................................... Pedilus granti, NEw SPECIES .............................. Pedilus 1810181 (Horn) ...................................... THE LABIATUS GROUP ........................................ Pedilus lonaflobus Fall ..................................... Pedilus crotch! (Horn) . . . .................................. Padilla labiatl‘s (Say). 0 o o 0 o e oooooo o oooooooooooooooooooooo .0 o Pedilus iomae, NEW SPECIES ........ . ..................... Pedilus monticolus (Horn) .................................. THE LU GU BRIS GROUP . . ..................................... Pedilus W18 (Say). . . .. ................................. Pedilus cyanipennis Bland .................................. THE TERMINALIS GROUP ..................................... Pedilus canaliculatus (LeConte) ............................. Pedilus impress” (Say) . . . ................................. Pedilus terminalis (Say) .................................... UNPLACED SPECIES .......................................... Pedilus serratus Fall . . . . . ................................. Pedflus alticolus Fall. . . ................................... 113 114 119 121 124 127 130 133 138 145 149 152 156 161 162 165 171 174 179 180 183 188 194 200 205 206 ~ 214 217 218 226 231 235 236 240 TABLE OF CONTENTS, continued INCERTAESEDIS......OOOOOOO ....... 0.... OOOOOOOOOOOOOOOOOOOO Corphyra calypso Wickham ........... . . ............ . ...... FIGURES ............................................. REFERENCES ................................................ APPENDIX: Plant Associations for North American Species of Pedilus 243 243 245 286 9. 10. ll. 12. 13. 14. 15. LIST OF TABLE Seasonal Distribution of North American Pedilus ,,,,,,,,,,,,, Cantharidin Associations for North American Pedilus AdultSOOOOOOOOOOO?OOOOOOOO ....... O 0000000000000000000000 Suprageneric Character - Character State List ............... Suprageneric Character Analysis ...... . . . .- ................. Characters and Character State Polarity Hypotheses for North American Species of Pedilus ......... . ............ Character State Distribution Matrix: Pedilus Species Groups ................................................. Characters and Character State Polarity Hypotheses for the punctulatus Species Group (1-9) and punctulatus Subgroup (lo-I3) ............................... Character State Distribution Matrix: punctulatus‘ Species Group (1-9) and punctulatus Subgroup (lo-13) .......... Characters and Character State Polarity Hypotheses for the lewisi Species Group ............................... Character State Distribution Matrix: lewisi Species Group........... ..... ........................... Characters and Character State Polarity Hypotheses for the labiatus Species Group. ............................. Character State Distribution Matrix: labiatus Species Group.............. ........ . ............. ......... Characters and Character State Polarity Hypotheses for the terminalis Species Group ........... . ............... Character State Distribution Matrix: terminalis Species Group ........................................... Geographical Distribution Matrix ........................... xi 44 45 53 54 57-58 59 62 63 66 67 70 71 78 LIST OF TABLE, continued 16. Pedilus inconspicuus (Horn): Analysis of Elytral Color Morphs in Western North America .......................... 142 17. Pedilus lugubris (Say): Analysis of Pronotal Color Morphs in Michigan .............. . ...................... 212 18. Pedilus lugubris (Say): Analysis of Pronotal Color Mal-pm in Eastern North America .......................... 213 19. Pedilus canaliculaws (LeConte): Analysis of "Red" and "Black" Pronotal Morphs in Two Michigan ' Localities ............................................... 224 20. Pedilus cmaliculatus (LeConte): Analysis of "Red“ and "Black" Pronotal Morphs in Northeastern North America OOOOOOOOOOOO O ..... O O OOOOOOOOOOOOOOOOOOOOOOOOOOOO 225 xii 2. 10. ll. 12. l3. 14. 15. 16. LIST OF PLATE North American Faunal Zones and Abbreviations. . . ...... . . . . . Pedilus spp. and Hadronema princeps Uhler at Cantharidin-baited Filter Paper Discs . . - . . . . . ...... . ........ Proposed Phylogeny for North American Pedilus speciwenumeeoooooeocooeoooooooeoooooooooooooooooooooo Proposed Phylogeny for Members of the punctulatus Species Group............. ............................... Proposed Phylogeny for Members of the lewisi Species Group...00.00....0............OOOOOOOOOOOOCOOOO ...... .0 Proposed Phylogeny for Members of the labiatus SPeCiQGroup 000-000.0000000000001‘000000000000 ........... Proposed Phylogeny for Members of the terminalis SpeciesGroup................ ...... ..... ....... Regions Referred to in Keys to Female Pedilus spp. . . . . . . ..... Pedilus Adult: General Anatomical Structures AssociatedWith the Head ........ PedilusAdult: Antennae PedillwAdmt: AntennanOOOOOO0.0.0.0...00.000.000.000... Pedilus Adult: General Anatomical Structures AssociatedWiththeThorax ..... . Pedilus Adult: General Anatomical Structures Associ- atedWiththeLe@00000000000oooo'ooooeoooooooooooooo ..... Pedilus Adult: General Anatomical Structures Associ- atedWiththeElytra................... ............ Pedilus Adult Females: General Anatomical Structures Associated With the Abdomen . . . . . . ....... . . . . . . . Pedilus Adult Males: General Anatomical Structures Associated With Abdominal Sections 9 and 10, and Pmmeres0.0......OOOOOOOOOOOOOOOOOOO0.0.000...O. ....... xiii 27 29 56 61 65 69 73 77 245 247 249 251 253 255 257 259 LIST OF PLATE (cautioned) 17. Pedilus Adult Males: General Anatomy of the » Parameres .............................................. 261 18. Pedilus Adult Males: General Anatomy of the Median Lobe for the flabellatus Species Group and punctulatus Species Group (in part) .................................... 263 19. Pedilus Adult Males: General Anatomy of the Median Lobe for the punctulatus Species Group (in part) and the lewisi Species Group .................................. 265 20. Pedilus Adult Males: General Anatomy of the Median Lobe for the labiatus, lugubris, and terminalis species groups for P. set-rams and P. alticolus ....................... 267 21. mewvaoooo 00000 o ooooooooooooooooooooooooooooooo 269 22. Pedilus lugubris (Say) Larva: General Anatomical Structures .............................................. 271 23. Geographical Distribution Maps for North American Pedilus: P. flabellatus (Horn), P. parvlcollis Fall, P. (11110er (Horn) .......................................... 273 24. Ggraphical Distribution Maps for North American P us: P. dentatus Abdullah, P. nisticus Abdullah, P. fenderi sp. nov. .......................................... 275 25. Geographical Distribution Maps for North American Pedilus: P. flexiventris Fall, P. cavatus Fall, P. johnsonorum 3p, now, P. incmspicuus (Ham) ................ 277 26. Geographical Distribution Maps for North American Pedilus: P. vittatus (Horn), P. oregonus Fall, P. bardi (Horn), P. MCLUIOLUS LeConte ............................ 279 27. Geographical Distribution Maps for North American Pedilus: P. picipennts Fall, P. slogans (Hentz), P. grantl sp. nov., P. lewisi Horn ............................. 281 28. G aphical Distribution Maps for North American Pedl : P. longilobis Fall, P. crotchi (Horn), P. labiatus (Say), P. joanae sp. nov., P. monticolus (Horn), P. lugubris (Say), P. cyanipennis Bland ................. 283 29. G aphical Distribution Maps for North American Pedi as: P. canaliculatus (LeConte), P. impressus (Say), P. terminalis (Say), P. serratus Fall, P. alticolus Fan ............................................ 285 xiv INTRODUCTION For the most part, species of Pedilus occupy temperate and montane areas of the Holarctic region. The Palearctic and Nearctic faunal components contain roughly the same number of species, although that of the former has never been formally reviewed. Species richness is greatest in western North America; twenty-one species are known to occur in California alone. I first became interested in the genus while collecting adult Coleoptera at flowering Crataegus in southern Michigan. In several areas, Pedilus was the most abundant insect, yet very little was known about the group. A cursory review of the literature revealed a number of problem areas which further stimulated my curiosity. Females were regarded by most authors as largely undeterminable and, in spite of a great deal of color variation in material collected, even from a single tree, most keys relied heavily upon color. I was also very intrigued by the fact that the larva of Pedilus had not been described. This paradox was aptly stated by Dury (1902): "At certain 'haw' trees when in blossom, by holding my umbrella inverted under the branches and striking them a hard blow with a stick, Corphyra [=Pedilud would shower down, hundreds to a tree, and yet the larvae are to me absolutely unknown, nor have I the slightest idea where to 100k for them." Although the North American species of Pedilus were revised in a series of papers by Abdullah in the 1960's, much confusion remained concerning species identification. Many of the species were defined on the basis of color characters which are sexually dimorphic and far too variable to be of value for discriminating between many sympatric species. Keys were also based largely upon males, thus making identification of female Pedilus all the more improbable. My interests in the genus Pedilus encompass the world fauna. Data on the North American species will continue to be gathered as efforts are expanded toward a revision of the Palearctic fauna and a synthesis of hypotheses relative to the bionomics, phylogeny and historical biogeography of Pedilus as a whole. BIONOIICS OF THE NORTH AMERICAN SPECIES OF Pcdflus Virtually nothing was known about the life history and habits of Pedilus when this study was initiated, aside from the fact that adults could be collected, often in large numbers, on flowers. Although the puzzle remains far from complete, enough pieces have been discovered to offer a hypothetical life history. Larvae Mamaev (1976) reported collecting an unidentified Pedilus larva from rotting walnut twigs during the month of June. In El Dorado County, California, larvae of P- inconspicuus (Horn) were observed in late April and early May feeding inside decaying acorns of 011970113 kelloggi Newberry (Wharton, 1979). Nine additional larval collections are known, three each from California and Michigan, and single collections from Arizona, Minnesota and Virginia. In addition, three adults of P- vittatus (Horn) were reared by W.C. Gagne (label data and in litt.) from roots of the common nettle, U rtica (110900 Linnaeus. Habitat data associated with the larval collections share several characteristics. All larvae were associated with dead vegetative materials, on or just beneath the soil surface, and in microhabitats which would approach 10096 relative humidity. Pedilus larvae are probably phytophagous or omnivorous, and fungi, which proliferate under such conditions, may play an important dietary role as well. Size differentials of larvae collected together and at different times of the year suggest at least four larval instars. One to several years may be required to complete larval development, as two instars appear to be present in two collections, and early instar larvae were collected in El Dorado County, California, during adult emergence of three species in late June. Pine Pupation in northeastern and western species generally occurs in April to early May, somewhat later in the higher elevations of the Appalachian, Rocky and Sierra-Nevada Mountains, somewhat earlier in southern and south-central California at lower elevations. The pupal stage is probably spent in the same habitat as that of the larva. The only specific information relative to the pupal stage of Pedilus comes from a single female lugubris reared by the author from a mature larva. The larva, which was maintained at room temperatures under laboratory conditions, pupated on 5 April 1977 and emerged 8 days later. During this time, the exarate pupa remained relatively inactive. Adllts SEASONAL DISTRIBUTION AND LONGEVITY. North American species of Pedilus appear to be univoltine and, as can be seen in Table l, are active primarily from April to July, although records span all but November, December and January. Average adult activity for a given Species is roughly 12 weeks; life span for a single adult appears to be 10-15 days based on one reared specimen of P. 11.19qu and numerous specimens of P. canaliculatus, P. elegans, P. impressus and p. lugubris which were collected in the field and maintained outside in cages as well as indoors under laboratory conditions. With the probable exception of P. flabellatus and P. parvicollis, which appear to be nocturnally or crepuscularly active, adults of North American Pedilus are diurnal. MATING. In spite of having observed and collected thousands of male and female pedflus at flowers and on foliage during the day and night, a copulating pair has never been seen, nor were any such references noted in the data associated with museum specimens examined. This, mating may take place apart from commonly visited plants, perhaps on the ground where females may oviposit. In view of the fact that most species are commonly found feeding at flowers, pollen and nectar may serve an important function in egg maturation, with mating occurring intermittently or toward the end of adult life. FLOWER ASSOCIATIONS. Field observations and bionomics data associated with museum specimens suggest that most species of North American Pedilus are non-specific as regards flower species frequented (see Appendix for plant associations or individual species for actual records). Rather than selecting flowers which are closely related in a phylogenetic sense, Pedilus adults usually visit flowers or inforescences which display an overall spherical or radiate symmetry (the haplomorphic and actinomorphic type classes, respectively, of Leppik, 1957). The beetle flowers or "cantharophiles" (Delphino, 1868-75) which Pedilus adults have been or might be expected to be associated with include such taxa as the Magnoliaceae, many Rosaceae, Hydrophyllaceae (taxa such as some Phacelia, which are functionally haplomorphic), Cornaceae, Umbelliferae, and Compositae (taxa such as Monolopia, which are functionally actinomorphic). CANTHARIDIN ORIENTATION. The adults of numerous North American Pedilus demonstrate a positive orientation toward cantharidin, the bicyclical monoterpene chemical defense mechanism of meloid and some oedemerid beetles (Table 2). Results of present field work with cantharidin bait and an extensive review of the literature, to be presented elsewhere (Young, in preparation), have revealed that several groups of insects orient to the compound, or to meloid and oedemerid beetles which are known to produce it. In several instances, the meloid/oedemerid is simply a food source and cantharidin is utilized as a means of finding the prey/host. Cantharidin orientation on the part of pedflus and several other groups of beetles somewhat closely related to the Meloidae and Oedemeridae remains a mystery. With few exceptions, all specimens of Pedilus observed at cantharidin bait have been males. Once in contact with baited filter papers, specimens become very active and male-male attempted copulation has been observed commonly. Individuals also work over the baited filter papers extensively with their labial and maxillary palpi and mandibles. Apparent feeding activity on the part of p. lugubris males "attracted" to the meloid, Meloe angusticollis: has been noted in the field, and similar observations were reported for p. joanae (Leech, 1934, under the name monticolus) and p. terminalis (Pinto and Selander, 1970). In each case, it was the meloid elytra which were noted to have been chewed. On several occasions, pedflus have been observed chewing on the elytral apices of other male Pedilus at the baits or in crowded containers, and a few museum specimens of male peqilus have been examined which had the elytral apices partially chewed away. If the apparent feeding behavior in response to cantharidin-baited filter papers is homologous to chewing on meloid elytra, and if chewing on elytra of other Pedilus is homologous to chewing on meloid elytra, intuitive reasoning would support the hypothesis that male pedilus also contain cantharidin or a cantharidin-like compound in the elytra. Following this notion, it is interesting to note that the highly modified male elytral organs of many species of North American Pedilus are undoubtedly sites of chemical production as supported by the smooth surface with numerous pores leading to an internal cavity (Figs. 52- 54). The admittedly speculative scenario presented above suggests that perhaps cantharidin or a closely related compound was present in the ancestral stock which eventually gave rise to Pedilus, meloids and perhaps oedemerids. NATURAL EN EMIES. Although there are few specific records, adult Pedilus are known to be attacked by tiger beetles (e.g., P- 91990713, p.170), ants (e.g., P. abnormis, p118, parasitic mites (e.g., P- 1001103, 9. 199 ), spiders (e.g., P. canaliculatus, p223), and lizards (e.g., P. inconspicuus, p143). PHORESY. Phoretic deuteronymphal mites have been recovered from adult specimens of P. joanae (see p. 199 ) and P. johnsonorum (see p.136). In each instance, the mites were congregated on the ventral surface of the beetle, primarily on the abdomen. 8 TABLE 1. Seasonal Distribution of North American Pedilus. Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec abnorm is alticolus bardi canaliculatus cavatus crotchi cyanipennis dentatus elegans f enderi flabellatus flexiventris granti impressus inconspicuus joanae johnsonorum labiatus lewisi longilobus lugubris monticolus oregonus Darvicollis Dicipennis Duncullatus "usticus 8erratus term inalis Vittatus \ TABLE 2. Cantharidin Associations for North American Pedilus Adults. Pedilus Species Nature of Association Source of Information P. abnormis P. bardi P. canaliculatus P. cavatus P. crotchi P. elegans P. impressus P. inconspicuus P. joanae P. johnsonorum P. labiatus P. lewisi P. longilobus P. lugubris P. monticolus P. oregonus P. picipennis P. punctulatus P. serratus P. terminalis b) a) b) a) b) a) b) a) b) b) b) c) at cantharidin bait on Eupompha slogans at cantharidin bait at cantharidin bait at cantharidin bait at cantharidin bait on Meloe niger at cantharidin bait on Meloe angusticollis at cantharidin bait at cantharidin bait on M eloe niger at cantharidin bait at cantharidin bait near caged Epicauta fabrici at cantharidin bait at cantharidin bait at cantharidin bait on Meloe angusticollis at cantharidin bait at cantharidin bait at cantharidin bait at cantharidin bait at cantharidin bait at cantharidin bait on Meloe americaruls on Meloe angusticollis at cantharidin bait New Record New Record New Record New Record New Record New Record Harrington (1894) New Record Say (1827) New Record New Record Leech (1934) New Record New Record Abdullah (1964b) New Record New Record New Record New Record New Record New Record New Record New Record New Record New Record Pinto 6: Selander (1970) Pinto 6c Selander (1970) New Record HISTORICAL REVIEW FOR NORTH AMERICA The genus Pedilus was erected by Fischer von Waldheim in 1822 to receive his fuscus from the Altai mountains of south-central Siberia; he attributed the new gems to the heteromerous family Pyrochroidae. Five years later, Say (1827) became the first individual to publish on the North American fauna when he described lugubris, collaris and terminalls (all from Missouri), labiatus (from "'Missouri Territory"), and impressus (from Pennsylvania). Although they were described under the generic name Anthicus, Say was obvi0Isly not satisfied with the designation: "[these]...differ much from the other species of the gems that have come under my observation; but as they appear to me to approach more closely to this gems than to any other, 1 place them here for the present." He later placed them in the genus Lagria (Say 1835) stating that their characters, "seem to justify, if not the formation of a new genus, certainly a division of the present, under the name of Corphyra." In 1830, Hentz described elegans and infumata, attributing them with some doubt to the Palearctic gems Pyrochroa. Newman (1838) was first to correctly assign members of our fauna to the genus Pedilus when he described fulvipes, imus, lugubris, guttula, and rufithorax. In the same year, Randall (1838) published his description in inornata from Maine, utilizing Pyrochroa as the generic name. Pennsylvania was listed by Ziegler (1844) as the type locality of his nigricans, haemorrhoidalis, mficollis, and marginicollis. Following the description of marginicollis, he added, "Var. a. Thorax entirely destitute of the thoracic spot." The first diagnostic key was provided by LeConte in 1847. At that time he also described one new species, P. pulcher from Kentucky and listed nigricans 10 11 Ziegler as a synonym of imus Newman, haemorrhoidalis Ziegler as a synonym of elegans (Hentz) and both rufithorax Newman and marginlcollis Ziegler as synonyms of infumatus (Hentz). Four years later LeConte (1851) described Pedilus punctulatus in the "Lagriariae" (= Lagriidae), perhaps in deference to Say's contribution of 1835. The type locality was listed as San Francisco, and this punctulatus became the first known species from the western states. In his synopsis of the "Pyrochroides" (= Pyrochroidae) of the United States, LeConte (1855) treated Pedilus just ahead of Pyrochroa. At this time, he stated that punctulatus was found on flowers during the month of June. He also added a number of names to the growing list of synonyms for the eastern species of Pedilus In LeConte's "Classification of the Coleoptera of North America" (1861-1862), Pedilus again shifted position-this time being treated as the only member of the "group" Pedili (Anthicidae:Pedi1ini). The next species to be \ described was cyanipennis. from Virginia (Bland, 1864). Jacquelin-Duval (1859- ! 1863) noted apparent differences between Corphyra and Pedilus stating that they differed in the structure of the abdomen. This usage was followed by LeConte (1867a) in his "List of the Coleoptera of North America," wherein Corphyra is treated as a member of the anthicid tribe Pedilini. During the same year, LeConte (1867b) also published a description of C. canaliculata, a new Species from Ohio. The western fauna became the foets of attention for roughly the next decade as Horn (1871, 1874, 1883) described ten new species from Colorado (C. lewisii) and California (vlttata, funebris, abnormis, crotchii, monticola, incon- spicua, bardii, distinguenda, and flabellata). In his 1871 paper, he stated that fl-lnebris and punctulata LeConte were found on composites. Of the eastern 12 impressus (Say) he stated, "I have lately seen specimens caught under circum- stances admitting of but little doubt in the supposition that impressus is merely the male of collaris." In his 1874 and 1883 contributions, Horn expressed doubts as to the validity of funebris, stating that it might be synonymous with punctulatus. In terms of bionomics, Horn (1874) noted that, "the species of Corphyra may be taken in considerable numbers, at times, on the flowers of various umbelliferous plants." He also alluded to the importance of the male genitalia in terms of identifying the species, and provided the first figures (1883). The relative position and description of the group was not changed in LeConte and Horn (1883), the only difference being in the replacement of the generic name Pedilus by that of Corphyra. The name change was not followed by Champion (18900), however, and Casey (1895) retained the name Pedilus when he \ spoke briefly of the "marvelously complex...intromittent apparatus" of the males. In his revised list of the beetle fauna of Cincinnati, Ohio, Dury (1902) speculated that pulchra and fulvipes were but synonyms of labiata and lugubris, respectively. He also reported collecting large numbers of canaliculata from blossoms of "white thorn" and buckeye. Blatchley (1910) provided brief diagnoses, habitat notes, and a key to the species of Corphyra known from Indiana. The contribution also contained a description of one new species, C. brunnea, and the statement was made that newmani was probably synonymous with fulvipes. In Pics (1911) contribution to the Coleopterorum Catalogus, all North American species of the taxon were attributed to Corphyra which was, in turn, treated as a subgenus of Pedilus. Wickham (1914) described C. calypso from the rich Miocene deposit at 13 Florissant, Colorado. This is the only known fossil representative of the genus. A total of ten western species was described under the generic name Pedilus by Fall in 1915, including arizonensis (Santa Catalina Mtns., Arizona), oregonus (Oregon), and infectus, longilobus, serratus, cavatus, flexiventris, alticola, lineatus, and picipennis (all from California); flavidus was also proposed "for the sake of convenience" as a color variation of inconspicuus. The characters associated with the male genitalia were utilized more than in any prior work, with Fall stating that he was unable to satisfactorily identify the species in any other way. In terms of bionomics, Fall wrote, "the beetles themselves are found- -sometimes in numbers—on flowers, appearing rather early in spring—March and April—in the vicinity of the Coast, but considerably later in the season in the Sierras." He described one additional species, parvicollis, from Tulare County, California, several years later (Fall, 1919). The "higher" classification of Pedilus became the subject of diSCIssion again in 1920 as Blair referred the taxon to the Pyrochroidae. During the same _ year, Lang (1920) published his well known catalog of North American Coleoptera in which Pedilus is treated as the only member of the tribe Pedilini, within the family Pedilidae; Corphyra is listed as a synonym of Pedilus. The resemblance of Pedilus to certain of the pyrochroid genera was alluded to by Van Dyke (1928) as he stated, "Pedilus parvicollis Fall, I would place as an undoubted Dendroides..the shape of the head, prothorax, and elytral features as well as the generic and family characters are those of the latter and not of Pedilus" Fall (1929) was quick to express his disagreement but added that "as a matter of fact there is no appreciable error in saying that reliable differences of family rank between Dendroides and Pedilus are non-existent." Crowson (1955) seemed 14 somewhat uncertain when he placed Pedilus and its relatives in the Anthicidae stating that, "according to Bbving and Craighead it is not possible to distinguish these groups in the larval state, though it should be recalled that they said the same of the larva of Anaspis, a gems certainly quite distinct from these forms in imaginal characteristics." In volume four of his series dealing with the beetle fauna of the Pacific Northwest, Hatch (1965) reiterated one of the unsolved taxonomic problems regarding Pedilus as he wrote, "I am unable at present to identify females except when taken with males..." In addition to a key to the males of ten regional species, his treatment included the description of one new species, pratti, from Washington, as well as an illustration of the male of P. cavatus'Fall. Abdullah (1962, l964b, l964d, l964f, 1966b, 1969) published on the North American species of Pedilus, dividing them into 14 "natural" taxa, based primarily upon the structure of the male genitalia. Four groups, including six new species, were treated in his first paper (1964b). The new additions were from California (nsticus, inurbanus, and dentatus), Kansas and Iowa (medias), Eastern United States (parvus), and New York (sulcatus). In the next paper (l964d), two more groups were considered, and flavidus Fall was elevated to species level. A new genus, Neapedilus, was erected by Abdullah in 1969 for parvicollis Fall, "...to be consistant with the classification of other tribes and subfamilies of Anthicidae." In 1976, Mamaev briefly described the larva of an unidentified species of Pedilus which he collected from rotting walnut twigs on the Sary-Chelek l'eservation, Kirgiz SSR. This was the first published account of the larval stage of the genus. Pedilus larvae were further characterized at the generic level by I"awivrence (197 7). Only quite recently, however, has the first species description 15 of a Pedilus larva been published. Wharton (1979) collected and reared larvae of P. inconspicuus (Horn) from decaying acorns in western El Dorado County, California. Doyen (197 9), relying in part upon the descriptions provided by Mamaev and Wharton, compared Pedilus and related heteromeroIs larvae to those of Cononotus METHODS Phylogenetic considerations at both the suprageneric and intrageneric levels are based largely upon morphological characters of the adults, since information on other stages, larvae in particular, is presently lacking for many taxa, as are data relative to cytogenetics, behavior, internal morphology, etc. The character complex associated with the male genitalia has played an \ increasingly significant role in the classification of North American Pedilus ever since Horn (1883) observed that "the form of the male sexual apparatus" could be utilized to separate species. The male genitalia have been referred to extensively in defining the species groups of Pedilus and in separating several closely related species which Show considerable intraspecific variation in other characters. Anatomical Terminology. Except as noted specifically in the text, terminology follows that in current use for the Heteromera (e.g. Lawrence, 1971 for larvae; Crowson, 1955 and Doyen, 1966 for adults). LARVAE Dorsal mandbular armature (Figures 138, 139). As noted by Lawrence (1 977:51), "Shiny patches or vaguelines on the basomesal portion of the dorsal 16 mandibular surface may be seen with light microscopy...these areas contain fine, cuticular elements, usually flattened, which are inclined and which face in a basomesal direction." Elma m and Ema! rods» These terms are utilized as described by Lawrence (1977:49): "The ventral part of the mouth cavity just above the labiomaxillary complex is strengthened by sclerotized hypostomal ridges, which may continue posteriorly as hypostomal rods. These rods [absent in Pedilud are usually divergent or subparallel and occur in many groups of Heteromera." Pa'abasal rim As defined by Young (l975:9), "well developed raised lines...on the dorsum of the abdominal segments just posterad of their anterior margins." These ridges also are found in association with the thoracic segments in some groups. \ Urogomphd plate. Young (l975:8) Ised this term in a broad sense to N encompass the entire ninth abdominal segment. It is restricted here to include only the highly modified ninth abdominal tergite. Urogunphal pits. This terminology follows Young (1975:8): "The two I deeply pigmented, heavily sclerotized excavations between the urogomphi." / These are the cul-de-sacs of Moody (1880), the fossae of Hayashi (1969b), the "pouch—like imaginations" (sensu Mamaev, 1976) and the intraurogomphal pits of Doyen (1979). ADULTS all-face awlpturagand Punctatlan. Although difficult to quantify, dorsal Surface sculpturing and microsculpturing of the head and pronotum are diagnostic for a number of species. Punctation of the head and pronotum is 17 referred to as coarse if the individual punctures are as large as or larger than the facets of the compound eyes, and dense if the punctures are generally separated by a distance less than their diameters. Since punctation associated with the head commonly varies according to particular subregions (e.g. clypeal, frontal, postocular), descriptions and keys for certain species are so referenced. Heteromeroid troclnnters (Figures 39—41). As figured by Crowson (1955: figure 106), the trochanter is obliquely attached to the base of the femur which is, in turn, acutely prolonged around the inner aspect of the trochanter. Basal tooth of tarsal claw. With the exception of the diagnostically elongate, truncate "tooth" exhibited by the males of longilobus (Figure 47), the basal tooth may be referred to as small and poorly developed (Figure 46) or well developed (Figures 42-45). Since each of these types is easily distinguishable, and a broad range of structural variation can be seen in the "well developed" type, it was unnecessary to further quantify the relative size of the basal tooth associated with the tarsal claw. Internal flame of metatharacic coxae (sensu Crowson, 1955: figures 151- 152) (Figure 41). A component of the endoskeleton which is visible externally as a groove of variable length along the surface of the coxa. Hylecoatotd type matanrbstarnite (Figure 38). This term was derived from the name of the lymexylid gems Hylecoetus According to Crowson (1938), "The Lymexylonid Hylecoetes has a furca that seems one of the most primitive in the order..." ‘ Elytral apices The hind angle of the elytron in adult Pedilus is normally rounded and slightly less than 90° (Figure 51). In several species, the angle is more acute (Figures 49-50), and in both the males and females of labiatus (Figure 18 48), the apex is conspicumsly acuminate and is referred to as dentiform. Males of bardi have the elytral apices not only conspicuously acuminate but also upcurved, a condition referred to as caudate Elytrul 0mm The males of many North American species of Pedilus possess variously modified organs in association with the subapical or apical regions of the elytra. These regions may be variotsly impressed and/ or swollen and are Isually impunctate and shining as seen with light microscopy or microscopically por0Is as seen with the scanning electron microscope (Figures 52-54). They are also commonly of a different color than the remainder of the elytron, although intraspecific color variation is sometimes great. As previOIsly hypothesized (p. 6), these organs may be involved in the synthesis of cantharidin or a cantharidin-like compound. Female Abdominal Terminalia The terminology Ised in this study is largely that of Tanner (1927). The ventral bacculus and oblique bacculus are sclerotized rod-like vestiges of valvifers one and two, respectively, while the lateral extension of. the ventral bacculus is synonymous with the baccular acumination of Ekis (1977:18). The coxite is a lobate, lightly sclerotized structure which Doyen (1966:140) referred to as "valvifer 2." Tanner observed that the coxites are secondarily divided in many groups. In Pedilus the coxite is obliquely divided into two coxital segments and the distal segment bears a setose coxital stylus (the gonostylus of Doyen, 1966:140). The dorsal and ventral laminae were described by Ekis (1977:18) as "membranous and acuminate laminae" which extend caudally "between the dorsal and ventral bases of the coxites." A lightly sclerotized lobe is located anterodorsad of the coxites. This is the proctiger of Tanner and may represent the tenth tergite. 19 Male Abdominal Terminalia. A highly modified structure of uncertain origin is located posterad of the eighth abdominal segment, enveloping the external genital apparatts. It probably involves elements of the ninth and tenth segments and aids in extrusion and retraction of the external genitalia. The sclerotized lateral portions of this structure are referred to as the ninth hemitergites (the spicule plates of Doyen, 1966:138); they fIse anteroventrally to form a heavily sclerotized rod-like apodeme, the spiculum gastrale. The dorsal sclerite between the ninth hemitergites is believed to represent the tenth tergite. Although Wood (1952), Lindroth (1957) and others have attempted to stabilize terminology applied to the external elements of the male genitalia in the Coleoptera, a cursory examination of current literature indicates that such a goal is far from a reality. The author recognizes some of the problems involved in establishing sound homologies and has deferred largely to the terminology of Sharp and Muir (1912), more for the sake of consistency than because it is believed to be preferable on morphological grounds. A As defined by Lindroth (1957:246), the tegmen is "the (single or double) sclerite situated basally (proximad) of the penis [= median lobe! and often surrounding it when in repose. The tegmen is Isually divided into basal piece and parameres." Basal piece, the sclerotized, unpaired proximal portion of the tegmen. Parameres, as defined by Lindroth (1957:244), "a pair of appendages (sometimes coalescent or even completely joined) forming the distal (apical) part of the tegmen and usually protruding on each side of the penis [= median lobd." These are the lateral lobes of Sharp and Muir (1912). 20 Median lobe, the penis as defined by Lindroth (1957:244): "The apical (distal), unpaired part of the copulatory apparatus, containing the terminal portion of the orifice of the ejaculatory duct." Orientation of tegmen, according to Crowson (1955), "The Heteromeroid type of aedeagus [= tegmen + median 1006 seems to have developed from the Cucujoid type by the disappearance of the ventral part of the ring-piece of the tegmen, leaving the param eres attached to a dorsal basal piece...ln several families of Heteromera this arrangement undergoes a further inversion (probably by a twisting of the end of the abdomen comparable to that occurring in male Diptera) with the final production of an apparently ventral trilobe type of tegmen..." For present purposes, the normal heteromeroid orientation is defined as tegmen located dorsad of the median lobe, while the inverted heteromeroid orientation characteristic of Pedilus is defined as tegmen located ventrad of the median lobe. Digiti laterales, articulated lobes associated with the parameres of many Pythinae and Salpinginae (Spilman, 1967:7). Ventral sinus (of the parameres), the excavated distoventral surface of the fused parameres, particularly well developed in joanae (after Ekis, 1977:18). Use of scanning electron microscopy has revealed a number of previously unknown structural components of the median lobe in Pedilus. The adjective "phallic" has been Ised in association with such structures in order to conform to the convention proposed for similar structures found in the clerid genus Perilypus (Ekis, 1977). Phallic papillae and phallic pectinations were seen along the distoventral surface of the median lobe in several species (Figures 115, 122, 131). They appear to be associated with the distal orifice of the ejaculatory duct and may aid in moving sperm out of the duct at the time of ejaculation. A well defined phallic cord was observed along the ventromesal surface of the median 21 lobe in lugubris (Figure 125) and cyanipennis (Figure 126). A pair of plate-like structures is associated with the distoventral aspect of the median lobe in a number of species of Pedilus (Figures 100-107, 109, 111- 115). The inner margin of each plate is conspicumsly serrated; they are referred to collectively as the paired serrated structures and may function analogms to the phallic plicae and phallic papillae of Perilypus as hypothesized by Ekis (1977: 18). Alix-aviation and Nomenclatlral Convention. Following each species description is a brief geographical distribution, a list of those collections which are known to have specimens, the total number of adult male and female specimens which were available for study, and peak seasonal flight time for adults. Geographical distribution data follow the format and abbreviations adopted for use in the U.S.D.A. Catalog of the Coleoptera of America North of Mexico (e.g. Atkins, 197 9) (Figure l). Abbreviatiors for collections are arranged according to the four-letter system developed by Arnett and Samuelson (1969); a list of the collections referred to in this study is provided in the "Acknowledgments" section. Maps depicting hypothetical species distributions (Figures 142-171) were prepared by extrapolation of verified geographical locations in conjunction with geomorphological considerations. Although these distributional ranges should be considered conservative for the most part, few substantial extensions are expected. In the interest of economy, only those species for which fewer than 75 specimens or 10 different collection records were available are considered "poorly represented." Detailed label data for the following poorly represented 22 species are listed under the species distribution: flabellatus, parvicollis, denta- tus, rusticus, crotchi, serratus, and alticolus However, date - locality and any additional pertinent label data were extracted from each specimen examined and are available upon request. These data are filed by species and cross referenced according to the collection from which they came. Several Pedilus epithets have been based on modern personal names of males (i.e. bardi, crotchi, fenderi, granti, lewisI). In spite of the grammatical considerations imposed by Isage of Latin, I concur with the objections raised by Spilman (1979) on the use of "-i" and "-ii" as stated in the International Code of Zoological Nomenclature, and agree: "The we of an -i ending regardless of the original ending is the better solution." Keys. The males of most species may by distinguished by secondary sexual dimorphisms associated with the antennae and elytral apices (i.e. the presence of elytral organs in addition to primary sexual characters associated with the genitalia. The key to males stresses all these characters; fortunately, males typically die with the tegmen at least partially extruded, thereby virtually eliminating the time consuming process of "pulling" genitalia prior to examina- tion. Species determination of females remains very difficult, and several of the western species are still unassociated. For this reason, three regional keys to the females have been prepared, thereby substantially reducing the number of species treated in any single key (see Table 15 and Figure 9). In order to maximize their usefulness to nonspecialist as well as specialist users, an attempt has been made to stress identification in these keys; evolutionary relationships are not implied. 23 Users should find that males are far easier to key out than females, the characters being far less subjective. Collecting Pedilusin series to better insure that at least 1-2 males are present is certainly recommended, but it should be cautioned that several species may commonly be found on the same plant or other habitat. Types. Data relating to type locality and repository are provided in the species synonymy which heads the discussion of each species. In many cases, additional observatiors relative to the examination of type material have been provided. These comments follow the synonymy and are headed "Type Informa- tion." Dissectilg Methoth. Males Isually die with the genitalia at least partially extruded. In most instances, enough can be seen to readily allow for species determination. However, in those instances where structural detail is obscured in the male or when female genitalia need to be examined, the following sequence may be followed for brittle specimens: detach the abdomen and place in a solution of warm potassium hydroxide (KOH) for 3-6 hours (note: if abdomen cannot readily be detached without risking damage to legs, etc., first place whole specimen in boiling water for 20-40 minutes to slightly relax); remove abdomen from KOH, rinse in acetic acid, rinse in 7096 ethyl alcohol (EtOH); pull genitalia and examine; store in genitalia vial with glycerine. In the case of males, the author frequently removed the genitalia from the abdomen and often separated the median lobe from the tegmen. However, to reduce the risk of losing track of small structures in a genitalia vial, it might be advisable as a general rule to leave the genitalia attached to the abdomen. [moments Size ranges of species in the adult stage were derived by measuring the smallest and largest specimens from the total series available for 24 study, independent of sex. While females tend to be, on an average, slightly larger than males, a great deal of overlap was noted and no informational content would be provided by listing size according to sex. Actual length is defined as the distance along the meson between the apex of the labrum and the distal end of the elytra in the case of adults, or the mesa] distance between the apex of the labrum and the distal end of urogomphi in the case of larvae. mistratiom. Rough drawings were made in pencil from an image projected by a Batsch and Lomb (#42-63-59) Microprojector, and corrected for' detail visually from observations made with a Wild MS-97049 stereoscopic microscope. Scanning electron micrograpls were made from a positive-negative Polaroid film (type 665) imaged through an International Scientific Instruments ([81) Super Mini Scanning Electron Microscope. Unless noted otherwise in the acknowledgments, all illustrations and S.E.M. photomicrographs were prepared by the author. Collectilg Techniques. Adult Pedilus may be collected, frequently in large numbers, from flowers of many angiosperms, particularly those belonging to Leppik's (1957) haplomorphic and actinomorphic "type classes" based on floral symmetry. Adult emergence appears to be synchronized with the flowering period of such taxa as Prwuls, Crataegus and Ribes in the east and western montane regions, and in the far west with Prunus, Ceanothus, Phacelia, Hera- cleum,and Conium Sweeping, beating vegetation and foliage, and malaise-trapping have all proven to be productive techniques as well. Meadows, particularly those near lakes or streams with mesic surroundings, provide the best habitats for utilizing such methods. Few species orient to light at night, but those which might be collected at light traps (i.e. flabellatus and potentially parvicollid are rarely encountered by any other means, 25 Several thousand specimens representing 20 of the 30 North American species of Pedilus have been collected at cantharidin bait and dead meloid beetles during the course of these studies. Baits (Figures 2-3) are prepared by dissolving cantharidin crystals in an organic solvent, such as acetone, and dipping filter papers into this solution. The acetone quickly evaporates, leaving the cantharidin impregnated between the fibers of the filter papers which are kept in covered plastic petri dishes. Specimens found in the petri dish may be collected by simply placing the lid back on the bait. Since the specimens are usually very active, they may be most easily killed by placing the dish, cantharidin and all, directly into a freezer. When recovering the baits, it is wise to check surrounding vegetation for additional specimens as they commonly land a‘ short distance from the bait and continue orientation behaviors while walking. These specimens can be collected by hand or with a modified bulb—type aspirator. (Note: avoid the me of a standard aspirator to prevent cantharidin from accidently being inhaled.) The baiting technique could easily be modified into a trap, thereby ‘ reducing the amount of monitoring involved. Specimens of Pedilus orienting to the bait eventually wander away, presumably as a result of short term sensory conditioning. 26 PLATE 1 Figure 1. North American Faunal Zones and Abbreviations. Figure 1 . UT A2 A2 Arlzona BC Brltlsh Columbla CA Calllornla CO Colorado CT Connecticut DC Olatrlct of Columbla OI Delaware F1. Florlda 0A Goorgla OL Greenland IA Iowa ID Idaho It. llllnols IN Indlana WY MOUNTAIN SOUTHWEST 27 CO MI Manltoba MD Maryland ME Malno MI Mlchlgan MN Mlnnaaota MO Mlsaourl M8 Mlsslsalppl MT Montana NI Now Brunswlclt NC North Carollna ND North Dakota NE Nebraska NF Newfoundland NH New Hampahlra NJ New Jersey NM New Mexico NS Nova Sootla NT Northwest Torrltorlos NV Nevada NY New York OH Ohlo OK Oklahoma 00‘- M \. utafls‘ ON Ontarlo OR Oregon PA Pannaylvanla PE Prlnca Edward Island PM St. PIorro-Mlouolon PO Quebec Ill Rhodolaland SC South Carollna 90 South Dakota 3K Saskatchewan TN Tennessee TX Texas UT Utah VA Vlrglnla VT Vermont WAWashIngton WI WIaconaln WVWoat Vlrglnla WY Wyomlng YT Yukon Tarrltory LAW \ 0 M M“ Figure 2. Figure 3. 28 PLATE 2 Pedilus abnormis (Horn) and the mirid bug, Haw-enema princeps Uhler, on cantharidin—baited filter papers in Wyoming. Pedilus canaliculatus (LeConte) at cantharidin bait in Michigan. 29 CRITERIA FOR SPEC- RECOGNITION AND FOR INTRASPECIFIC AND SUPRASPECIFIC GROUPINGS Mayr's (1969:412) "biological species" definition is insensitive to such considerations as asexual modes of reproduction and paleontology. As a theoretical basis for recognition of species, the following modification of Simpson's (e.g. 1961:153) and Wiley's (1978:18) "evolutionary species" definition has been adopted: A species is a single lineage of ancestral-descendant populations of organisms which maintains its genetic identity from other such Engages and which has its own evolutionary tendencies and historical The species included in this study, whether redescribed or proposed for the first time, are operational hypotheses of what are believed to be the "biological" or "evolutionary" species of North American Pedilus. Pragmatically, however, application of the above definition to taxonomic endeavors—particularly revisionary studies such as this which involve a number of species—is rarely successful in view of our lack of knowledge in the area of population dynamics and our inability to determine the extent of gene flow between populations. As Erwin (1970) pointed out, "When only miseum specimens are utilized, different criteria are necessary to supplant those given in the ...[theoretical species definition] ." For the purposes of this study, the author concurs with the criteria listed by Wheeler (197 9): Sympatric or a110patric populations which differ in at least one consistent genitalic character are considered two distinct species; allopatric populations which differ in one or more consistent characters, not attributable to the normal range of intraspecific variation, are considered two distinct species. 30 31 This is not to dispute the numerous examples cited by such authors as Btsh (1975) and White (1978) for which such criteria might lack sensitivity. Certainly, different modes of speciation—particularly those involving parapatric and sympatric models—have given rise to many taxa for which other criteria might be required to resolve genetically isolated sympatric or parapatric species which maylack morphometric differences in genitalic structure. Until the Palearctic faunal element of Pedilus can be studied and until more data are available relative to larvae, supraspecific phylogenetic relationships will be discussed in terms of informal species groups. The six provisional species groups of North American Pedilus are formed on the basis of synapomorphy, with hypotheses of relative plesiomorphy and apomorphy determined largely by in-group and out-group comparisons, correlation with character transformation series, and Dollo's principle regarding the general irreversibility of evolutionary processes. GENERIC AND SUPRAGENERIC PHYLOGENETIC CONSDERATIONS As previmsly mentioned, Fischer von Waldheim (1822) attributed Pedilus to the family Pyrochroidae. This familial association was followed by LeConte (1855) and Blair (1920). The segregation of Pedilus along with several other genera into the family Pedilidae is credited to Lacordaire (1859). He envisioned two groups ("Pédilides vrais" and "Scraptiides") as comprising the family and listed N ematoplus LeConte under Pédilides and Tanarthrus LeConte under Scraptiides as "genre incerte sedis." The genera making up Lacordaire's "'I‘ribu Scraptiides" are currently assigned to the families Euglenidae (Xylophilus Curtis), Scraptiides (Scraptia Latreille and Trotomma Kiesenwetter) and Anthicidae (Tanarthms LeConte), while N ematoplus LeConte is now generally attributed to the Cephaloidae (Abdullah, 1965b; Mamaev, 1973; Lawrence, 1977). This leaves Lacordaire's Pedilidae with Pedilus, Eurygenius LaFerté, Stereopalpus LaFerté, Macratria Newman, and Ster0pes Steven. Although a number of genera related to Eurygenius and Stereopalpus have been described since 1859, this is essentially the familial association followed by Pic (1911), Leng (1920), Arnett (1960-1963), and Lawrence (1977). A third scheme of classification has been to associate Pedilus with the Anthicidae (sensu lato). Such an association was implied by Say (1827) when he described the first North American species of Pedilus under the generic name Anthicus Paykull. LeConte (1861-1862, 1867a), LeConte and Horn (1883), Crowson (1955) and Abdullah (1962-1969) have supported this broad view of the Anthicidae. 32 33 In brief comments relative to the phylogeny of Heteromera, Crowson (1966:512) envisioned four lines evolving from an ancestral stock, "with larval characters much like the existing Zopheridae, and adults more similar to Synchroa and Stenotrachelus" "From such an ancestor, we might derive (l) the Aderid-Anthicid-Meloid line (2) a line leading via Pythidae and Pyrochroidae to Salpingidae, Mycteridae, Boridae and Inopeplidae (3) one leading via Synchroid and Zopherid-like forms to Merycidae and Monommidae and Colydiidae, also perhaps to true Zopheridae and the Tenebrionid group of families (4) a line to Melandryidae and Mordellidae-Rhipiphoridae, also to Scraptiidae." In attempting to evaluate Crowson's scheme relative to the Pedilidae (auctorum), one cannot help but wonder whether the "Aderid—Anthicid-Meloid line" really is separable in a phylogenetic sense from the "line leading via Pythidae and Pyrochroidae to Salpingidae, Mycteridae, Boridae and Inopeplidae." Thus, in entertaining the question of where to place Pedilus, adults of a number of taxa representing both of these "lines" have been examined as well as one additional group, the Oedemeridae, which Crowson (I966) "intentionally omitted from discussion" (Tables 3-4). Since larval stages of nearly half of the taxa under consideration are either unknown or inadequately described, characters of larvae have not been included in the analysis. A few comments on interfam ilial relationships based upon larvae will be offered following a discussion of the character analysis based upon the adults. l. 5. 7. 34 Agatha (AGNA). This monotypic genus is based upon the European decoratus Germar, a single male of which was examined. An'motria (ANIS). In the course of examining Pedilus taken at cantharidin bait, several males of an undescribed genus and species collected by Mr. Gary Shook and Mr. Albert Allen were discovered. The taxon, which is being described elsewhere (Young, in press), is considered at this time in view of its close relationship to Pedilus. Anthicidae (ARTE). I prefer to follow a somewhat narrower view of this family than that proposed by Crowson (1955) and further expanded upon by Abdullah. Species examined: N otoxus spp., Lagriomorpha indigacea Young, Lemodes sp. Relevant data have also been included from Chandler (1977a; 1977b) and Werner (1964). . Boridae (BORE). A small family whose members have frequently been referred to the Pythidae. The character analysis is based upon specimens of Boros unicolor Say. Cmtus (CONO). A single C. macer Horn was examined. Copobaerm (COPO). This genus, with two species in Chile and two recorded from Argentina, was originally placed in the Pedilidae (Fairmaire and Germain, 1863) but has recently been associated with the Anthicidae, sensu lato (Abdullah, 1969). A single specimen of C. tristts Fairmaire and Germain was examined and compared with observations reported by Abdullah (1969:334-335). Buygeniinae (BURY). Specimens examined: Retocomus marinas (Halde- man), Pergetus campanulatus (LeConte), Stereopalpus mellyi LaFerté, S. vestitus (Say), Bactrocerus concolor LeConte, Rilettius rahmani Abdullah, 10. 35 and Leptoremus argenteus Casey. Additional sources of data included the following: Pergetus Casey (Abdullah, 1960), Leptoremus Casey (Abdullah, 1961a), Egestria Pascoe (Abdullah, 1961b; 1969), Duboisius Abdullah (Abdullah, 1961c; 1964c), Bactrocerus LeConte (Abdullah, 1963a), Pseudo- bactrocerus Abdullah (Abdullah, 1963a), Neoeurygenius Abdullah (Abdullah, 1963b, 196411), Eurygenius LaFerté (Abdullah, 196411), Cadogenius Heller (Abdullah, l964g), Qadrius Abdullah (Abdullah, l964i), Rilettius Abdullah (Abdullah, l964i), and Stereopalpus LaFerté (Abdullah, l965d; 1965e). Ictistygnini (ICTI). Material of this group was not available for examination; the character analysis is based upon Borchmann (1936) and Abdullah (1969). mcolloganius (INCO). This genus was erected by Pic (1916) to receive his testaceipennis of Madagascar. Abdullah (1965f) considered the genus a primitive pyrochroid. I am reluctant to accept Abdullah's broad interpretation of the Pyrochroidae, but having not had an opportunity to examine any specimens of Incollogenius it will not be associated with a particular family at this time. Data used in the character analysis have been extracted from Abdullah (1965f). WMGD). Fairmaire and Germain (1863) referred this aberrant genus to the family Lagriidae; it has since been associated with the Melandryidae (Champion, 1890a), the Oedemeridae (Champion, 1916) and the Anthicidae (Abdullah and Abdullah, 1968; Abdullah, 1974). The character analysis is based upon specimens of L. australisChampion and L. obscurellaFairmaire and Germain which were available for examination, as well as a brief generic description provided by Abdullah and Abdullah (1968). ll. 12. 13. l4. 15. 16. 36 lea-atria (MACE). Characters utilized in the following analysis are based upon observations of M. marina (Fabricius) and M. brunnea Casey. Meloidae (HELD). The meloids constitute a rather well defined group compared to most other heteromerous taxa. However, if one examines Protomeloe Abdullah, the distinctions become less sharply defined. Specimens of Lytta sayi LeConte were examined in addition to the literature descriptions involving Protomeloe (Abdullah, 19641; 1965g). Mitreelabus (mm). The author has had to rely upon Abdullah's (1969) description of this small (1-2 species) Chilean genus. Mycteridae (MYCT). Specimens of Mycterus scaber Haldeman, M. canescens Horn and Lacconotus sp. were examined, and data relative to Phaeogala Fairmaire were extracted from Abdullah (1965c). Oedemeridae (OEDM). As noted by Crowson (1955:135), "This family seems to be relatively clear-cut and natural, but its precise affinities are far from clear." Two species which are considered by many to represent the least specialized lines were selected for critical examination: C 0100113 augustus LeConte and Nacerdes melanura (Linnaeus). Otlniidae (OTHN). For present purposes, a narrow definition of this family will be followed. No material of Trogocryptus or related genera was available for study, and no attempt has been made to critically examine the aberrant genus Aegialites, an othniid (sensu Crowson, 1955) or a salpingld (sensu Spilman, 1967). In this study, Elacatis umbrosus (LeConte) was examined. 17. 18. 19. 20. 21. 37 Pyroclroidae (PYRO). As understood here, the family is restricted to the Pyrochroinae of Young (1975) and Lawrence (1977). Larvae of Ischalia Pascoe have now been positively identified, thus necessitating the removal of the Ischaliinae (serial Young, 1975). Lawrence (1977:43) referred Lemodes Boheman to the Pyrochroidae, but this genus appears to be closely related to the aberrant anthicid genus Lagriomorpha Champion, as noted by Young (1978:108). Species examined: Schizotus cervicalis Newman and Pyrochroa coccinea (Linnaeus). Pythidae (PYTH). The character analysis is based upon specimens of Pytho americanus Kirby and Trimitomerus riversi Horn. Although Young (1976) has demonstrated that Sphalma quadricollis Horn is a pythid, it has been excluded from consideration. The adult exhibits several probable autoapomorphies which would make the analysis far less homogeneous. Sta-apes (STIR). Specimens of Steropes caspius Steven were examined to augment Abdullah's (1966a) observations. Techmeminae (TECH). Although a close relationship between Techmessa Bates and its allies and others such as Pilipalpus Fairmaire and Cycloderus Solier has been commented upon by a number of coleopterists, Paulus (1971:84) was apparently first to formally recognize them under one subfamily. He attributed the assemblage to the Pyrochroidae, in deference to Crowson's broad view. Specimens of Techmessa sp. were examined while data on Pilipalpus and others were extracted from Abdullah (l964c; 1965a; 1965f; 1967). Trictenotomidae (TRCT). Members of this small family appear to be rather closely related to the Pythidae; Trictenotoma grayi Smith was examined in this study. 38 The 21 taxa referred to above are contrasted in the character analysis presented in Table 4. This is, admittedly, a crude analysis. The characters chosen reflect, for the most part, those commonly cited relative to one or another group of Heteromera. And, while certain commonly referred to I characters (e.g., "type of antennae") which appear to be of little value in ascertaining interfamilial relationships have been excluded, many of the 17 characters selected could well have polyphyletic origins. For this reason, assignment of plesiomorphic and apomorphic character states has been attempted for only a few of the 17 characters utilized, and these are offered as but tentative hypotheses. In spite of these obvious shortcomings, a number of relationships may be pointed out. If the digiti laterales (sensu Spilman, 1967:?) associated with the parameres ' of the male genitalia of BORI, PYTl-I, TECH, and TRCT can be considered as homologous structures, out group comparisons would support the hypothesis that the character is apomorphic, thereby bringing these four taxa into close relationship on the basis of synapomorphy. Other groups with digiti laterales include the AegIalitinae (Spilman, 1967) and Salpinginae, Dacoderidae, Inopeplidae, and Trogocryptinae (Lawrence, 1977). If Trogocryptinae is correctly placed in the Othniidae, the lack of digiti laterales in Elacatis would have to be considered secondary (i.e. apomorphic within the Othniidae), and thus quite unlike the plesiomorphic absence of the structures in other Heteromera. Abdullah (1965f) and Paulus (1971) have pointed to a relationship between [H CO and TECH: indeed, a superficial resemblance is inescapable. The only significant differences between the two groups involve the male genitalia, with 39 INCO lacking the digit! laterales and having the parameres free as opposed to the fused condition observed in TECH. However, only a few specimens of Techmessa have been examined and further comments will be withheld pending the study of additional techmessine genera. Crowson (1955:135) commented, "A notable structural feature of the Anthicids is the great shortening of the internal flange of the hind coxae..." An in group comparison of Heteromera suggests that the "reduced" character state is apomorphic, thereby indicating a close relationship among ANTH, COPO, EURY, ICTI, LAGD, MITR, and STER. Other characters which lend support to this association include externally open and internally closed prothoracic coxal cavities (ANTH, COPO, most EURY, and LAGD), and parameres fused throughout their entire length (ANTH excluding Notoxinae, coro, EURY, lCTI, LAGD, and MITR). Abdullah (1969:353) and Lawrence (1977:44) treated MACR as a subfamily of ANTI-l. Although the base of the internal flange of the metathoracic coxae is not reduced in MACR and the parameres are free distally, it does exhibit externally open and internally closed prothoracic coxal cavities as well as several additional characters common to ANTH (e.g. normal heteromeroid orientation of the tegmen and a closed radial cell). Certainly, the placement of MACR near ANTH is no more objectionable than suggesting a close phylogenetic tie between STER and AN TH. The absence of digiti laterales in association With the parameres 0f Pedilus precludes any direct association of its members' with BORI, PYTH, TECH or TRCT. Furthermore, the lack of evidence to support a secondary loss of the structures would omit OTHN and INCO from consideration as well. In Pedilus, the base of the internal flange of the metathoracic coxae is elongate, the 40 prothoracic coxal cavities are open externally and internally, and the parameres are free distally. This combination places a serious strain on any scheme of classification which would demand a direct relationship between Pedilus and ANTH, COPO, EURY, ICTI, LAGD, MACR, MITR or STER. As previome mentioned (p.15 ), the only groups of organisms known to produce cantharidin are MELO and OEDM. This observation, together with the fact that positive cantharidin orientation has been demonstrated in ANIS, ANTH, and PYRO in addition to Pedilus, leads me to suggest that cantharidin utiliza- tion/orientation has profound phylogenetic implications. While the current data relative to cantharidin may be quite incomplete, a crude hypothesis may be offered at this time: Cantharidin, or a cantharidin-like compound, was present in the ancestral stock which gave rise to ANIS, ANTH, MELO, OEDM, PYRO and Pedilus. A cursory look at the life history of MELO and a number of OEDM suggests that cantharidin has evolved in these two groups as a chemical defense mechanism. And, although the utilization of cantharidin in the remaining taxa is open to question, the preponderance of males observed at cantharidin bait suggests that the compound may be associated with reproductive behaviors. Imperative to this hypothesis is the idea that cantharidin production is apomor- phic. An examination of this premise by out group testing reveals that members of only one other family of beetles, the Endomychidae, are known to orient positively toward cantharidin. It is possible that cantharidin, or 'a similar compound, has been independently evolved in this group; it seems equally plausible that cantharidin may mimic some secondary fungal compound utilized by endomychids in host selection. In either case, the supposition that cantharidin production in the Heteromera is apomorphic would appear to be parsimonious With present data. 41 Since Pedilus departs considerably from MELO and OEDM in terms of cantharidin utilization (in numerals other characters, as well), and the lack of a direct relationship to ANTH has been demonstrated, potential affinities are limited to ANIS and PYRO. With respect to the characters examined, Pedilus differs from ANIS only in conformation of the compound eyes and, to some extent, in the structure of the tarsal claws. While a number of other characters, including antennal and elytral structure as well as shape of the pronotum, clearly point to generic recognition of ANIS, this taxon is, nonetheless, very closely related to Pedilus- The differences between Pedilus and PYRO appear to be equally slight. In Pedilus, the radial cell is closed and tegmen orientation is of the inverted type (although observations indicate that subsequent torsion may not be at all uncommon), while all the genera of PYRO examined possess an open radial cell and normal heteromeroid orientation of the tegmen. The two taxa also differ slightly in the structure of the tarsal claws. In addition to Pedilus. larval stages for 13 of the 21 taxa analyzed above have been at least partially characterized: ANTH, AGNA, BORI, CONO, EURY, LAGD, MELO, MYCT, OEDM, 0THN, PYRO, PYTH and TRCT. The highly specialized larval forms of MELO obscure phylogenetic relationships and will not be considered here. Of the known OEDM larvae, only those of Calopus show a relationship to other taxa under consideration. In fact, Rozen (1960) stated, "There is little similarity between the oedemerines and calopodines, and the calopodines share more characters with the larvae of families often considered to be related to the Oedemeridae...than do the larvae of either nacerdines or the oedemerines." 42 The larvae of BORI were described by St. George (1931, 1940); he considered them most closely related to PYTH. And, although the description of TRCT provided by Gahan (1908) is of little value by current standards, and the figures are highly stylized, this work would seem to support a relationship between TRCT and PYTH. Crowson and Viedma (1964) and Lawrence (unpublished mantscript) have critically examined the larval stages of MYCT, indicating a close relationship between its constituents and BORI. Crowson and Viedma also stated (p. 106) that, "the larvae of Lagrioida...is very unlike those of Mycteridae, but suggests a possible affinity to the Anthicid group...." Kitayama (1978) stated, "The presence in anthicids of the penicillus at the base of the mandibles is unique among the cucujoids. In combination with the latter and with the presence of one pair of ocelli and of sensory appendices on the antennae, the family, based on larval characteristics forms a natural group." However, other heteromerms taxa with but a single pair of stemmata are known, including 0THN (Hayashi, 1969a) and Scraptiidae (Hayashi, 1962) among others. And, as Doyen (1979) pointed out, the structure of the sensory appendix is similar among AGNA, ANTH, CONO, BURY, PYRO and Pedilus. Thus, only the presence of a mandibular penicillus appears to be apomorphic in ANTH. Based upon his analysis of larvae representing AGNA, ANTH, CONO, EURY, MYCT, PYRO, Pedilus and Salpingidae, Doyen (1979) concluded, "Pergetus [= EURYJ shares most characters with Anthicidae..." Although a mandibular penicillts is not present in EURY, the group remains very poorly known in terms of larval stages and a close relationship to ANTH should not be dismissed solely on this basis. The structure of the urogomphal plate, asperities associated with the 9th 43 sternite and spine-like setae associated with the legs support a relationship between 0THN and PYTH. The presence of 1-2 urogomphal pits on the caudal margin of the urogomphal plate, between the urogomphi, may be of great phylogenetic importance as suggested by Mamaev (1976:98): It is essential to point out that the pouch-like invaginations between the urogomphi are intricate formations, the presence or absence of which is of fundamental importance, since it may be used to converge a whole group of families—Pyrochroidae, Pythidae, Boridae and certain others. The greatest similarity in the structure of these formations is found in larvae of Pedilus sp. and Pogonocerus thoracicus Fisch.- W...of the family Pyrochroidae. Of the taxa under present consideration, urogomphal pits are present in AGN A, BORI, CONO, PYRO, PYTH, and Pedilus. On the basis of the above remarks, I submit that Lacordaire's concept of Pedilidae is untenable as is the broad definition of Anthicidae. Of the taxa examined, Pedilus appears to be most closely related to ANIS, both of which come close to PYRO. While the adults of both AGNA and CONO each exhibit a number of probable autapomorphies, characters associated with the larvae tend to support placement of these aberrant taxa between PYTH and Pedilus-PYRO. Either Pedilidae should be redefined to include only ANIS, Pedilus and perhaps AGNA and CONO, or PYRO should be expanded to encompass this entire ‘ assemblage. For the present, the latter scheme would appear to better illustrate the relationships indicated above. Likewise, EURY should either be elevated to family rank or included with ANTH. Until more is known about the larval stages of EURY and some of the aberrant genera of ANTH, the disposition of EURY must remain somewhat uncertain. However, the evidence indicates that EURY _ is certainly more closely related to ANTH than to the PYTH - PYRO - Pedilus assemblage. 44 on Any “mom Adv any mmasuouma Huwwmm.msfiumon mommasusm .na sumsma oufiusw wanna Apv “zaasumfiv ooum Adv nbv mmuoasuma mo wusuoauum .oH Heusuwa Adv “vauum>sa ADV “wwoumaououon Hmaho: Adv 00v smawou mo sowumuaofiuo .mH sausaso 0: any "was Amy Ifiuum zammum Ham moufisuoum Hmaaaovns manfimfi> .qa AwaHB usomnm Adv mumsuuov zauoom Anv "vsaamou Hams Amy oaomuonuwumav Mumam Hauunsonam mo ouaumz .MH 0: Anv amok Amy usomwum Awsws ofiosuonuwuoav Hams mmvoz .NH 0: Any «mom Asv vomoau Awe“: ofiumuonusuoav Haws Heaven .HH sundae Ifivcmmmm Ho uuoao Adv “nuoou Human Anv mmHaaHm Amy mSMHu Hummus we make .OH oumaauflaama was ousaauasaonwuss Adv "musaauassoa ADV “manage Amy mucoawom wonoa “Hausa mo coaumEHOMaoo .m msxoo vmusvou Anv “mummaoam Amv ofiomHOSuMuoa mo owsmam Hmsuouaw mo swam .w assumumomoa mo vmmoaouwuss Awsfiumoa o: ADV “mum Adv sauna: HOV wawuwoa maumumaaw owowuonuomoz .m Hanan Adv momuo>mamuu AAV "wafiuUMmoum was Hmowaoo Amy mmxoo ofiusuosuoun mo oaks .0 on any mush Amy some haasauouaw moaufi>mo stoo ofiosuonuoum .m o: ADV «mos Amy some haasduouxm mmaua>so meoo uaosuosuoum .q on Anv «mom Amv haasHMusH noswmuma.a=uoaoum .m oussfiwuuam any mouwuau va some vssoqaoo «0 show .N on Any new» Adv :xoms: wswauom .mmho usanmn vouuauumsoo snow .H moumum Houomussu nouowumso .umHA mudum Hmuumumno I Houomusso uauofiummumsm .m wands 17 16 ab 15 as 14 13 abc 12 ab ab 11 ab ab 10 ab ab Character ab ab a a a Suprageneric Character Analysis Table 4. Taxon INCOLLOGENIUS LAGRIOIDA ANISOTRIA ANTHICIDAE BORIDAE cononorus COPOBAENUS EURYGENIINAE ICTISTYGNINI MACRATRIA’ MELOIDAE MITRAELABRUS AGNATHUS be ab MYCTERIDAE ab ab b? ab ab ab ab ab be be ab ab b TRICTENOTOMIDAE TECHMESSINAE PYROCHROIDAE OEDEMERIDAE OTHNIIDAE PEDILUS PYTHIDAE STEROPES INTRAGENEBIC PHYLOGENB'HC CONSERA'HONS The general inability to comment with reasonable confidence relative to out group character polarity among the Heteromera has been alluded to previously. The paucity of knowledge at this level together with the fact that a significant Palearctic Pedilus fauna remains virtually unstudied prohibits the formulation of any detailed phylogenetic hypotheses at this time as regards the North American Pedilus fauna. Nevertheless, a number of remarks can be made to indicate Mble evolutionary relationships among the species umder consideration. In the first of his papers dealing with the taxonomy of Pedilus, Abdullah (1964b) divided the gems into 14 species groups which he considered "natural" or "monophyletic, at least so in a broad sense." While characters associated with the male genitalia are certainly of great phylogenetic significance, my interpretation of these and other characters forces me to conclude that most of Abdullah's 14 "natural" groups are, in fact, paraphyletic or polyphyletic. For this reason, it would be less confusing to entirely disregard Abdullah's informal groups as opposed to making necessary changes and attempting at the same time to preserve the names Dr. Abdullah applied to them. Thus, the species group names proposed herein bear no direct relationship to the names Abdullah (l964b— 1969) referred to, although some conceptual overlap is inescapable. Table 5 provides a listing of characters useful in ascertaining the intrageneric relationships of the North American Pedilus fauna. Character state polarity hypotheses have been derived largely from in group (= North American Pedilus spp.) comparisons, although a cursory examination of several Russian 46 47 species has certainly entered into the decision process. On the basis of these hypotheses, six species groups are recognized which accommodate 28 of the 30 species under consideration; two species (alticolus Fall and serratus Fall) are treated as unplaced species for the time being. From the character state distribution matrix (Table 6), a phylogeny of the six North American Pedilus species groups is proposed (Figure 4). Several interpretations seem plausible in attempting to formulate polarity hypotheses for character states associated with the basal tooth of each tarsal claw (Table 7, character 5; Table 9, character 4). The argument might be made that a small tooth forms a transformation series between no tooth, the condition found in most Pyrochroinae, and a well developed tooth. Were this the case, a small tooth would be considered plesiomorphic and a large basal tooth, apomorphic. In supporting such an hypothesis, one would have to accept the idea that a large basal tooth was independently evolved at least three times to account for the character state in P. abnormis and P. dentatus of the punctulatus group, in three of four species belonging to the lewisi group, and in the lineage which gave rise to the labiatus, lugubris and terminalis species groups (Figure 4). On the other hand, since the "well developed tooth" character state is prwent in at least some members of five out of six North American Pedilus species groups, this state might be viewed as plesiomorphic and any subsequent reductions as apomorphic. In following this logic, one would have to accept at least three independent reductions in the size of the basal tooth: once in the lineage giving rise to the flabellatus species group, once within the punctulatus species group and a third time to account for the character state observed in P. picipennis of the lewisi species group. This interpretation would also fail to 48 provide a logical means for deriving the "well developed tooth" character state from the "no tooth" condition seen in most Pyrochroinae. If, however, the "small tooth" character state was found in the ancestral stock and the "large tooth" state was evolved in the lineage which gave rise to the lewisi, labiatus, lugubris and terminalis species groups, only two independent evolutionary events would be required: one to account for the "large tooth" condition found in P. abnormis and P. dentatus, and a second to explain the "small tooth" of P. picu‘pennis. Since this explanation requires the fewest evolutionary steps and also provides a transformation series between the type of claw found in the Pyrochroinae and the types seen in Pedilus, it was utilized for polarity hypotheses in this study. Thus, the occurence of a small basal tooth is plesiomorphic in the flabellatus and punctulatus species groups and apomorphic in the lewisi Species group. The structural complexity of the elytral organ (see also p. 18 ) found in many North American species of Pedilus indicates that the character was probably derived early in the evolution of the group and has a common source as opposed to the notion that such a complex was independently evolved by members of several different lineages. Thus, in both the punctulatus (Table 7, character 6) and lewisi (Table 9, character 5) species groups the presence of this organ is considered plesiomorphic. Its absence in abnormis, inconspicuus, johnsonorum, rusticus, vittatus and picu’pennu’s is attributed to subsequent reduction. In adult females of all but two species of North American Pedilus, the lateral extension of each ventral bacculus is well developed and rod-like (Figure 63). Thus, the lack of a well developed lateral extension (Figure 64) in vittatus 49 (Table 7, character 13) and elegans (Table 9, character 7) is accounted for on the basis of secondary reduction and is considered apom orphic. The punctulatus group (Tables 7, 8; Figure 5) is defined by the paired serrated structures which are associated with the posteroventral surface of the median lobe (Table 5, character 6). In addition to the unique modification of the metathoracic tibiae in the male (Table 7, character 4), 1’. abnormis exhibits a number of other autapomorphies as well. The overall similarity in parameral structure between P. dentatus and the unplaced P. serratus is worthy of note, suggesting that serratus may represent an early off-shoot of the punctulatus group which subsequently lost the paired serrated appendages of the median lobe. Additional material of these rare species would certainly be useful in exploring this possibility further. The remaining species differ from P. abnormis and P. dentatus with respect to the deve10pment of the tooth associated with the base of each tarsal claw (Table 7, character 5). Two closely related assemblages of species are identifiable within the remaining members of the punctulatus species group: the cavatus subgroup and the punctulatus subgroup. P. cavatus and P. flexiventru’s represent closely related sister species within the cavatus subgroup. P. johnsonorum and P. inconspicuus cannot be placed in either subgroup, but are certainly more closely related to the punctulatus subgroup. The flabellatus group may be characterized by flabellate antennae in the male (Table 5, character 7), coarsely faceted eyes (Table 5, character 8) and by having the pronotum widest anteriorly. The first two apomorphies are commonly associated with nocturnally active groups, and indeed label data confirm this to some extent in the case of P. flabellatus Should such an activity pattern be 50 demonstrated for the exceedingly rare P. parvicollis, it would indicate a significant departure from the remaining species of Pedilus which are typically diurnal. The two species comprising this group are certainly sister species. Although evolutionary relationships between the North American Pedilus fauna and that of the Palearctic region cannot be commented upon at this time, it might be pointed out that the parameral structure of the flabellatus group is far more similar to that of several species examined from southern Russia than to any other North American species group. The similarity is particularly evident between P. parvicollis (Figure 69) and P. tibialis (Figure 66) or P. pallidipennis (Figure 67). The four species comprising the lewisi group (Tables 9, 10; Figure 6) are synapomorphous with respect to the excurvate parameral apex (Table 5, character 2) and the presence of a spine or tooth-like process associated with the ventral or ventrointerior aspect of each paramere (Table 5, character 4) (Figures 82-85). P. picipennu's exhibits a number of autapomorphies which set it apart from the remaining species (Table 9, characters 2, 3, 4, 5). Similarly, P. elegans is autapomorphic as regards antennal type (Table 9, character 1) and reduction of the lateral extension of the ventral bacculus (Table 9, character 7) and occurs only east of the Rocky Mountains, whereas the other three species are found only from the Rocky Mountains to the West Coast. P. granti is an isolated and perhaps relatively recent offshoot of the lewisi lineage, suggesting that lewisi or its immediate ancestors might have once occupied a much broader area of the extreme Southwest. If present interpretations regarding the placement of the labiatus species group are correct (Tables 11, 12; Figure 7), its characterization is more a 51 function of divergence by the lugubris-terminalis lineage than by autopomorphy. Wheeler (1979:262) noted a similar problem in the leiodid beetles and suggested that recent speciation might account for such a slight divergence. P. longilobus is autapomorphic with respect to the highly specialized truncate lobe of the tarsal claws in the male (Table 11, character 4). The delicately pectinate antennae of male P. crotchi (Table 11, character 2) set it apart from all other members of the group; similar variations manifest in the flabellate antennae of the flabellatus group, and the very strongly serrate antennae of P. serratus must be considered convergent, not a transformation series within a monophyletic lineage. P. labiatus, the sister group of the joanae- monticolus lineage, is autapomorphic as regards the dentiform elytral apices of both sexes. P. joanae and P. monticolus are sister species. The lugubris group is characterized by the dorsoventrally flattened para- meres (Table 5, character 1). Two eastern species, P. lugubris and P. cyanipen- nis, comprise this small group which is the sister group of the terminalis species group. Members of the terminalis group (Tables l3, 14; Figure 8) may be defined by their short median lobe which has the sides strongly convergent distally (Table 5, character 5) (Figures 127-129). Two distinct subgroups are recognized, each of which has several autapomorphies. The canaliculatus subgroup contains only P. canaliculatus and would appear to be the sister group of the terminalis subgroup which contains two sister species, P. impressus and P. terminalis. All three species are restricted to the northeastern states, being recorded no further south than North Carolina (canaliculatus and terminalis) and Missouri (impres- sus), and no further west than Kansas (impressus), Missouri (terminalis) and Illinois (canoliculatus). 52 The terminalis and lugubris species groups are the only groups restricted to eastern North America. Of the two unplaced species, possible evolutionary relationships for P. serratus were alluded to under discussion of the punctulatus group. The gross structure of the tegmen in P. alticolus bears some resemblance to that of P. abnormis, but the likeness may be symplesiomorphic. The spinulae associated with the inner parameral margins of P. alticolus represent an autapomorphy approached by no other Species of Pedilus. This is one of the "rarest" species of North American Pedilus based upon material available for study; the discovery and study of a permanent breeding population could add much to our knowledge of its evolutionary history. 53 sauoauoooe onoefis "m ooe>o u< auooeouo «o oeeem .m woumomm haomumou um wouoosm haosfim u< momm .w AaHumH mouawfimv oumHHoAsHm "m Amw ousmamv oumaauooq "a flow ouawwmv mumpuom samcouum no Ammnnm nouowflmv Buowaafim hauwoa .oumuummnam um mumuumm u< Aha mmhu amasoun< .5 Ah» onoa ceases mo commune Hmuuso> saga woumwoommm AmHHuHHH .moH .noHuooH nouamfimv usomouo "m usomne u< mousuoauun museums causes .0 Am~auhmfl nonemesv hafleo aHHeonHe umfiv usmwum>aoo hHwaouum macaw .uuonm um wouoamu mmvfim .muamsoao "4 nbv onoa amawoa mo mamsm .m Ammumm mmuawfimv sumsoa macaw hosivaa mmououn oxwa Aha ousasusn some unuoou muses .voaoao>on Hams m wawumon ”m oaaafim n< we assume uoaumuaaouuao> .q Ammuww mauswamv Aamumo mouswamv nbv amuoawuma voumumnom haaouums .haamsvsuw "m wousuomom sauces .hauasuns u< mo soaumusaom Houu=o> .m Amwumw mmu=Mfimv mus>h=oxw hamusus um ous>u=uxu haou=os uos u< any ouoasusa nuns mo nuns .N Aum .He mousmfimv hHHmuuso>omuov someounaoo maamuusm>ocuou vodouusam eamsoaowmmsoo "m haunwwam ou Hmuwuvawamunam u< any mouoasusm mo macaw .H ADV oasnuofiofi‘ on owning—Samoan Houooudno muoum Houomusno msawvom mo moaooam amofiusa< nuuoz How momwnuonhm huwumaom oumum umuooumso was muouooumno .m manna 54 Table 6. Character State Distribution Matrix: Pedilus Species Groups Character Species Group _ 1 2 3 4 5 6 7 8 9 flabellatus A A A A A A E B B punctulatus A A A A A B A A A lewisi A B B B A A A,B A A labiatus A A B A A A A,D A A lugubris B A B A A A B A A terminalis A A B A B A B A A u 55 PLATE 3 Figure 4. Proposed phylogeny for North American Pedilus species groups. 56 w_._u=oou oxwaunoax nua3 .3ouus: "m AHCH ouswamv sumuqusm xaasmuov Kama .hHHsumav UUHoQMu “a “nod ouswfimv Noam umuswoa saucaan wcaauom .haawumqv mumswasom no AcHHIHHH .ooH mouswfimv Aaoa ouswfimv :uOOu uaHoa usaan .vmvcsou ha Havana uaoum wswummn .eaamumav venuesu um umuaus ou mHHmumfiv wouoamu u< onoa cmavoa was: .w haamuucs> vmxoammv sawsouum "m Hashes n< muficumum Hmaaaovnm Sun was: .5 assess an ucomoua u< compo Hmuuhao mas: .o anomwuaov .voaoao>ws Has: “a uosaumwucw ou Hamam u< swag Humans mo :uoou Human .n moawusmamm usumfi3u hauswfiam s sanwu mnu msa>ww mafia .afiwuma uoauwumon sum: lance zaosvuano as madness .vossuumam wommusm yucca "museums maunwwam "m Hwauoa u< mamas caucuozumuoa was: .q mucuocan hammcov .eflomumoo no oumuoaan aaomcov .haomumoo sawumumvoa "m musuussa maomumem .haoafim u< coauMuossm Hauoaoum .m ouwuoaam sawmsov .mawmuwoo no masseuse hammcmv .haomumoo eaousumvoa um oumuocsa hammuwam .haoawm u< coaumuoaaa swam .N ouswsoao ouswsoam scuom .umamv um hkusuovoE amuse .mumuovoa.u< Axouuo>v was: we ousuaumo> .H AOV ownauoaon< on oasau036amsam uuuosusso ouwum wouonumno Amauoav asouwnsm mausasuocsm was Amuav macaw mowooam mausasuoanmnonu you momosuoaem hufiusaom oumum wouosusnu use muouosussu .n oHnsa 58 voosvmu hamsouum awn ouswfihv omsuno savanna: um>uso was canvass eoeoooed :er voaoam>ov Hams AHmums mousmfimv ooeoe Adamo: Adamo: 3V coamamuxo umasoomn Hmuucm> Hmumusq obv speeches mo samuma Ronda so nuoou Hmowamnsm xmam chumao was: souuham mass Hmuoucaouman .MH .NH .HH .OH ADV ounnuoaoa< A3 3.18286on ouuum Houosusso Houoousno A.v.u:00v .N mHan 59 Table 8. Character State Distribution Matrix: punctulatus Species Group (1-9) and punctulatus Subgroup (10-13) Character Species l 2 3 4 5 6 7 8 9 10 11 12 abnormis A A A B B B A A A bardi ' A C C A A A A A A A B A cavatus B A A A A A B E A dentatus A A A A B A A D A fenderi A A A A A A B E A flexiventris B B A A A A B ' E A inconspicuus A B A A A B A B A johnsonorum A B B A A B A C A oregonus A c c A A A A A A B‘ A A punctulatus A C C A A A A A A A A A rusticus A C B A A B B E B vittatus A C C A A B A A A A A B 60 PLATE 4 Figure 5. Proposed phylogeny for members of the punctulatus species group. 61 3.23055 .ucan mscouoco @332; mazofimcoofi Escocomccfi @3330 mtEono: tones. maoznac 233:2. 2.59:? 11 1O 6.12.13 8 3/ PUNCTULA 3,9 W TUS SUBGROUP CAVATUS SUBGROUP Figure 5. 62 voosvou mawdouum oocmusm assuouafi so eaaocwwunansm magmauc .Hacam scoops Hausa ououuaaa mammamu was zaomumoo samusuovoa masseuse haemaov was eaomusou haoumucvoa anomaaaw sauna: .susuummnsm wonoao>ov Hams Hmaamuma .voaoam>wv HH03 uaommua apomausov .voaoao>ov HH03 mumuoasn haomuunm .maoaam masseuse maomucmm .haoaam Quwhhmm A my scamsouxm “aqueous Hauuco> kuquA .5 Abv mmououa 33H unuoou no scans Hmuoamusm .o ammuo Hmuuhao was: .m 3nHu Hmmumu mo suoou Human .o coausuuaan Heuoaoum .m soaumuossa use: .N msdsouam mo mama .H ADV cannuoaon< 8V owsauoaoqmoam Houusucnu flufium HUUOQHQSU nacho moaoonm “manna ecu How mumsnuonam knapsaom ousum HQUUGHQSU fiflfl QHUUOQHQSU om OHDQH 63 Table 10. Character State Distribution Matrix: lewisi Species Group- Character Species 1 2 3 4 5 6 7 elegans B A A A A A B granti A A A A A B A lewisi A A A A A A A picipennis A B B B B A A 64 PLATE 5 Figure 6. Proposed phylogeny for members of the lewisi species group. 65 .330. ..cmco memos? 253203 1,7 2.3.4.5 Figure 6. 66 voaoam>uv HH03 Awe suswfimv mmoo Iona anomwuauv s ouaw vowaoaoua Ame muswfimv waomuw Beau osu ou numsoa ca Hansonam .onoa ousoaauu hHHsumav ousmsoao cm msaauom .vovasmxm ousuusam eaomsmv vac hflomucoo haoumuovoa Amm ouswwmv ousnwuoom oumuoaan kaomamv use aflomuooo kaoumumvoa usmmnm Hashes snowflussv .wonoam>ov Hams oumuuaan haomusam .haosam AeNImN mouswamv ousuumm ououoasm sameness .haosaw .4. Ahoy ADV 0.393803. ADV canau050Hmmam musum Houosusso macaw mowoonm enumaaoa saw you momosuonhm huaumaom oumum Houomumsu was muouususno msumauuma mo madam Hmuuso> .o oceans Hmuumam .m 9.0 v 330 Hmmuwu mo nuoou Human .q aOfiuuuoasm Hauoaoum .m 90v 093 1535. .N coausuossa use: .H Hmuosusno .HH wanna 67 Table 12. Character State Distribution Matrix: labiatus Species Group Character Species 1 2 3 4 5 6 crotchi B B B A A A oanae B A B A A B labiatus B A A A B A longilobus A A A B A A monticolus B A B ' A A A 68 PLATE 6 Figure 7. Proposed phylogeny for members of the labiatus species group. 69 9:002:08 oacso_ «320s. .co~oco o:ao:a:0_ :uQu 41 Figure 7. 7O AaNH ooewnmv woum>moxo hamaoaoaamaoo .msonann um Amwwuwma mouswamv manage as Amuv sped asauoa mo Nsn< .w Ab V mmououm Ammuqa mousmamv ussmoue "m assess u< mxHHunuoou Houmamusm .n Ana ouswfihv vo>uso havusssa um Ammucm amuswfimv unwanuum u< Auuv mouoaswsn mo xen< .o msouoaoosoo "m maouoaooan n< suueao was: .m musuuasa ousuoaan mascusnm hammusoo ”m haomaov .haomusoo maoumuovoa.u< nowusuossa Hmuuham .c Aem ousmwmv m=H=UfiHmaso HusvauwmaOH wawuson "n Hmauod u¢ abuoaoun we come: .m Aqm ouamamV consumaaomouoaa “m oumuosan sameness .Naoswm “4 soaumuoasa Hauocoum .N onSuaHsomouoaa um oumuoasn sameness .haosaw u< sowusuoasa use: .H AOV owseuoaon< on cannuoaowmoam Houooumno ousum Houosuono nsouo muduonm muasawauou man How oomunuonhm muauoaom macaw monocumno was muouosusso .na canmfi 71 Table 14. Character State Distribution Matrix: terminalis Species Group Character Species 1 2 3 4 5 6 7 canaliculatus B B B A A B A impressus A A A B B A B terminalis A A A B A A B 72 PLATE 7 Figure 8. Proposed phylogeny for members of the terminalis species group. 1.2.3.6.8 4.7 23583. 73 3393:: 2:230:23". Figure 8. HISTORICAL BIOGEOGRAPHY Certainly, no attempt to formulate testable biogeographical hypotheses can be made until the Palearctic Pedilus fauna has been examined. However, a few comments are in order at this time. Although admittedly somewhat artificial and not strictly conforming to physiographic constraints, the map provided for regional keys to adult female Pedilus (Figure 9) may be referred to in the following discussion. Continental North America has been divided into three faunal regions (Figure 9) which approximate "west of the Rocky Mountains" (Region 1), "the Rocky Mountains" (Region 2), and "east of the Rocky Mountains" (Region 3). Distributions of extant North American Pedilus species support such a division, suggesting that this physical barrier has played an important role in the biogeographical history of Pedilus, P. joanae being the only species to occur in all three regions. From the distribution matrix (Table 15), it can be seen that the greatest species richness (six species groups and subgroups and 20 species) occurs west of the Rockies. This suggests that western North America may have been the source of diversification for the North American Pedilus fauna, with subsequent faunal movement eastward. Supportive evidence for this scenario comes from the phylogenetic relationships of Pedilus species groups proposed above (Figure 4), wherein the strictly eastern lineage (lugubris and terminalis species groups) was hypothesized to be most closely linked to the labiatus species group which, in turn, is of western origin. The remarkable similarity in parameral structure between several Russian species of Pedilus and the two species of the flabellatus species group has 74 75 already been noted. These and other structural similarities support an hypothesis that the Palearctic faunal element is quite closely related to the pumctulatus- flabellatus lineage. A cursory look at several of the Palearctic species indicates that the western North American faunal element exhibits far more structural diversity. Thus, it is postulated that western North America served as the primary site for diversification of Pedilus as a whole, with two major subsequent faunal movements: one to the east which ultimately gave rise to the species groups of eastern North America, and the other to the north and thence west via the Bering land bridge which served to stock the Palearctic region. 76 PLATE 8 Figure 9. Geographical regions referred to in keys to female Pedilus spp. 77 Figure 9. Table 15. Geographical Distribution Matrix 78 Region 1 Region 2 Region 3 flabellatus group flabellatus parvicollis punctulatus group .abnormis dentatus rusticus fenderi flexiventris cavatus johnsonorum inconspicuus vittatus oregonus bardi punctulatus lewisi group picipennis elegans granti lewisi labiatus group longilobus crotchi labiatus joanae monticolus lugubris group lugubris cyanipennis terminalis group canaliculatus impressus terminalis unplaced species serratus alticolus KN NNNNNMNNMXMN N CLAMPICATION OF NORTH AMERICAN SPECIB Genus Pedilus Fischer von Waldheim Pedilus Fischer von Waldheim, 1822: 35. Type species, by monotypy, Pedilus fuscus Fischer von Waldheim. Corphyra Say, 1835: 189. Type species, by present designation, Anthicus lugubris Say. [Corphyra was originally proposed as a subgenus of Lagria Fabricius: Pic (1911) treated it as a subgenus of Pedilus Fischer von Waldheim. See further comments under "Remarks"! Neopedilus Abdullah, 1969: 358. Type species, by original designation, Pedilus parvicollis F all. NEW SYNONYI. 'Daaiption With general characters of Coleoptera: Polyphaga: Cucujoidea. ADULT Heteromerous: maxillae 2-lobed; tarsal formula 5-5-4 in both sexes; prothoracic coxae conical, prominent and projecting, trochanters of heteromer- oid type (sensu Crowson, 1955: figure 106); wings with 4 veins between Cubitus and jugal area (anal veins sensu Crowson, 1955: 91; anal + CU2 sensu Ponomarenko, 197 3: 454); metendosternite with narrow stalk and anterior tendons arising from the furcal arms: abdomen with 7 pairs of spiracles; male 79 80 genitalia of inverted heteromeroid type, with tegmen (= parameres + basal piece) oriented ventrad of median lobe; Malpighian tubules cryptonephridic. Head: Cranium (Figures 10-12) abruptly constricted behind eyes, forming broad "neck"; frontoclypeal suture present; labrum distinct, transversely rectan— gulate, fringed along anterior margin with short setae, tormal sclerite bifurcate, comprised of medial tormal process and lateral tormal process. Antennae (Figuures 18-30) ll-segmented, subfiliform to flabellate, usually serrate, inserted on frons between base of mandibles and emarginate compound eyes. Mandibles (Figures 14-15) subsymmetrical, conspicuously flattened dorsoventrally, wedge- shaped, apices simple to tridentate, adoral mandibular surface bearing densely setose, fleshy, lobe-like prostheca mesally and well developed, heavily sclero- tized molar surface basally. Maxillae (Figure 16), (Williams, 1938: Figure 41) with cardo large, produced basally into inner cardal process and outer cardal process, articulating distally with triangular basistipes; mediostipes arising from distal adoral aspect of basistipes. continuous with lacinia, basigalea small, subtriangular, giving rise to bulbous distigalea, apex of distigalea and adoral surface of lacinia densely tufted with moderately elongate setae. Maxillary palpus consisting of four setose articles arising from membranous apex of palpifer, article one small, article 2 slightly longer than 3, 4 cultriform and slightly longer than 2. Labium (Figure 17), (Williams, 1938: Figure 41): Submentum confluent with gular region; mentum distinct. trapezoidal; ligula broad, strongly emarginate, forming 2 fleshy lobe's, apex densely covered with short setae; labial palpus 3-segmented, article one small, 3 slightly longer than 2. Tentorium (Figure 13): Posterior tentorial arms connected at back of head by broad, transverse bar, the posterior tentorial bridge, and about halfway along 81 their length by narrow anterior tentorial bridge: anterior tentorial arms simple, gradually narrowed posteriorly to point of fusion with posterior tentorial arms. Thorax: Prothorax (Figures 31-33) with notum well developed, convex, sides rounded and lacking lateral margins, margined basally, weakly margined anteri- orly, slightly broader than head, usually ovate and widest at middle (wider anteriorly in several species): prosternal process acute; prothoracic coxal cavi- ties (Figure 32) completely open externally and internally, outer angles prolonged slightly, exposing trochantins. Mesothorax (Figure 36) with scutum transverse, anterior angles acutely prolonged anteriorly; mesothoracic scutellum small, shield-like; mesothoracic coxal cavities not closed outwardly by sterna; meso- thoracic episterna contiguous or nearly so anterad of mesosternum. Metathorax with ventral endoskeleton, or metendosternite, of hylecoetoid type (sensu Crowson, 1938, 1944) with slender stalk, well developed laminae and anterior tendons arising from furcal arms (Figuure 38). Legs (Figures 39-41) ambulatorial; internal keel of metathoracic coxae elongate; tibial spurs short, simple; tarsi with penultimate segment lobed below; pretarsi with pair of claws (Figures 42- 47), each typically produced into a well developed basal tooth (tooth reduced in several species, prolonged into large, distally truncate lobe in longilobus). Mesothoracic wings, the elytra, well developed in both sexes, covering thorax, except for pronotum and mesothoracic scutellum, and entire abdomen; surface irregularly, typically densely, somewhat rugosely punctate (moderately. sparsely and coarsely punctate in a few species, occasionally rugose), covered with short, retrorsely semierect setae: elytral apices variously modified in males of most species, impressed and/or conspicuously swollen, impunctate and devoid of setae, microscopically porous (Figures 52-54). Metathoracic wings (Figure 37) well 82 developed with radial cell narrowly open or closed, closed wedge cell. subcubital fleck present but poorly defined. Abdomen: Tergites 1-2 essentially lost, 3-6 poorly sclerotized, 7-8 setose and moderately well sclerotized. Female with 5, male with 6 visible, freely articulated sternites; sternite one absent, sternite 2 visible only as lightly sclerotized transverse band situated in membrane dorsad of metathoracic coxae: sternites 3-7 ( 99 ) or 3-8 (0'0' ) sclerotized, setose; hind margin of last visible sternite in female (= 7th) entire. often acutely rounded, that of male (= 8th) usually emarginate. Female Abdominal Terminalia: ’ Eighth abdominal segment typically retracted within 7th; 8th tergite (Figure 55) with hind margin entire or slightly emarginate; 8th sternite (Figure 56) giving rise to anteriorly projecting, heavily sclerotized, rod-like spiculum ventrale. connected posteriorly to ovipositor by telescoping membranous sheath. Ovipositor (Figures 62-64), (Tanner, 1927: Figures 72-73) with first and second valvifers reduced to paired, heavily sclerotized, rod—like oblique bacculi and ventral bacculi, respectively: each ventral bacculus giving rise along distal fourth of its length to well developed, rod-like lateral extension (strongly reduced in two species); distal ends of ventral bacculi articulating with inner anterior ends of oblique bacculi: proctiger colspicuously setose; coxites 2-segmented, basal segment much larger than distal; coxital stylus short, stout, setose, arising obliquely from distal coxital segment: dorsal and ventral laminae (sensuEkis, 1977: 18) distally acuminate. Male Abdominal Terminalia: Sclerites derived from abdominal segments 9- 10 (Figure 65) usually at least partially retracted within 8th abdominal segment; 83 9th sternite produced ventroanteriorly, forming heavily sclerotized, rod-like spiculum gastrale; 9th hemitergites contiguous ventroanteriorly, divergent ven- troposteriorly; distal margin of tergite 10 broadly emarginate, fringed with moderately elongate setae. Tegmen ventral, consisting of well developed basal piece broadly joined to parameres distally; parameres fused proximally, separate and subparallel to divergent distally; median lobe elongate. dorsoventrally flattened, produced basally into two apodemes, the median struts (sensu Sharp and Muuir, 1912: 482). LARVA While an attempt has been made to present a composite of species in the description which follows, it should be pointed out that only a few species are known at this time; the analysis is based largely upon the larva of Pedilus lugubris (Say). Mature larvae (Figures 133-134) attain lengths of 9-12 mm and widths of 1.5-2 mm. Body subcylindrical anteriorly and cylindrical posteriorly, its sides subparallel. Head and body lightly sclerotized, nearly white to light creamy yellow except for reddish-brown areas of heavy sclerotization such as mandibles, urogomphal apices, calli and urogomphal pits (sensuu Young, 1975: 8); relative mesal lengths of head (from apex of labrum to occiput) to thorax to abdomen approximately 1:2:6. Vestiture sparse, consisting of elongate dorsolateral setae , with a few shorter setae dorsally and ventrally. Head: Cranium prognathous, exserted from prothorax, vertex slightly flattened; moderately sclerotized and yellowish in color, lacking any definitive microsculpturing; vestiture consisting of a few elongate setae. Epicranial suture 84 with frontal arms lyriform, diverging anteriorly and extending to just behind antennal insertions; stem lacking, frontal arms thus confluent posteriorly with well sclerotized dorsomesal rim of occipital foramen; endocarina absent. Front- oclypeal region flattened; frontoclypeal suture lacking, but weakly impressed, mesally discontinuous transverse sulcus usually discernable between mandibular articulations. Anterolateral aspects of head with four (rarely three) pigmented stemmata on each side directly posterad Of antennal insertions, arranged as in Figure 134. Antennae prominent, about 0.4x as long as maximum cranial width, 3-segmented, third segment the shortest (l:l:0.5) and narrowest (1:0.8:0.4); second segment bearing a broad sensory appendix which is about 0.6)! the length and 2X the width of segment 3 and ventrad to it. Mouthparts supported posteroventrally by hypostomal ridges; hypostomal rods lacking; gula nearly as broad posteriorly as its mesal width, strongly narrowed anteriorly. Labrum symmetrical, width at base about 1.5X mesal length; epipharynx (Figure 137) with three stout setae along each anterolateral margin, one pair of marginal peg- like setae mesally, a pair of stout posteriorly directed setae mesally, flanked laterally by fine mesally directed setae; base of epipharynx bearing a dense cluster of short, posteromesally directed hairs, bordered laterally by numerous transverse, shingled plates which continue posteriorly along the pharyngeal wall to esophageal lining. Mandibles slightly asym metrical, large, heavily sclerotized and wedge-shaped, with triangular base and three apical teeth. Molar region of left mandible bearing a prominent distal tooth (Figure 138). that of right mandible lacking a tooth, bearing much coarser serrulations. Dorsal mandibular armature (sensu Lawrence, 1977: 51) well developed on both mandibles, with microcuticular combs oriented basomesally (Figure 139). Maxilla (Figure 140) 85 movable, free to base of mentum, composed of: l-segmented cardo, diagonally creased upward toward stipes, thus appearing 2-segmented; lightly sclerotized, undivided pad-like maxillary articulating area; maxillary male; and 3-segmented palpus. Mala shallowly cleft at apex, bearing many stout setae, especially along ventroadoral and‘apical aspects. Labium (Figure 141) with submentum trape- zoidal and longer than wide, mentum slightly longer than wide, broadest distally, prementum subrectangular, well developed ligula projecting anteromesally beyond palpi and bearing two setae apically; palpi 2-segmented. Hypopharyngeal sclerome (Figure 141) subrectanguular, anterior surface shallowly sulcate, poster- olateral angles bearing patches of setae. Anterior aspect of hypopharynx bearing numerous blunt, fleshy denticles laterally and apically, and a transverse row of setae just anterad of the hypopharyngeal sclerome. Thorax: Thoracic segments subequal in length. Prothorax with tergum, cervicosternum and basisternum somewhat more darkly pigmented than remainder of thorax; lacking parabasal ridge (Young. 1975: 9); cervicosternum well developed, divided into 3 plates; coxae large and subglobular; intercoxal distance (measured relative to coxal diameter) 1.25 diameters. Mesothorax and metathorax with terga very lightly pigmented, each with parabasal ridge developed; mesothoracic laterotergite bearing annular spiracle with accessory chambers, arranged as in Figure 135. Legs well developed, about 0.5x as long as width of mesothorax, each bearing a number of fine setae; coxa large and conical; femur and tibia subequal in length; tarsungulus well sclerotized and pigmented. Abdomen: Abdominal terga 1-8 and 9 (excluding urogomphi) subequal in length; parabasal ridge well developed on terga 8 and 9, inconspicuous on remaining segments; spiracular diameter about 0.7X diameter of mesothoracic 86 spiracle, annular with accessory peripheral air tubes directed posteriorly or posteroventrally (Figure 136). Ninth tergum (Figure 133) expanded posteroven- trally to form the entire terminal segment or urogomphal plate (sensu Young, 197 5: 8); bearing paired, heavily sclerotized, fixed, 2-branched urogomphi with inner branch curved mesally, outer (main) branch curved sharply upward at apex; two deeply pigmented, heavily sclerotized urogomphal pits well developed caudomesally, their surface variously microsculptured; pits narrowly separated by mesal ridge which is often produced into a blunt mesal tooth. Surface of urogomphal plate bearing a number of setiferous calli. Ninth sternum trapezoidal, somewhat recessed into shallow emargination of eighth sternum, bearing a single, stout callosity on each anterolateral aspect; tenth segment greatly reduced, visible ventrally as a narrow, poorly sclerotized sclerite, directed posterad of anus; usually bearing four setae, two on either side of meson. Distribution The Nearctic faunal element occurs from the southern portion of southern Canadian provinces south to the Gulf coastal states in the East and to the Mexican boarder in the far West, and from the East Coast to the West Coast. Species richness is greatest in California and lowest in the southeastern states. Diagnosis Adult - Head abruptly constricted behind eyes, thus forming a "neck"; eyes emarginate. Pronotum lacking lateral margins; prothoracic coxal cavities open externally and internally; prothoracic coxae prominent, conical and projecting; 87 tarsal formula 5-5-4 in both sexes; internal keel of metathoracic coxae elongate; tarsi with penultimate segnent lobed below; tarsal claws usually bearing a well developed basal tooth; elytra covering abdomen; males of most species with elytral apices variously impressed and/or swollen; radial cell closed; wedge cell closed; subcubital fleck present, poorly defined. Abdomen with all visible sternites freely articulated; males with tegnen of inverted heteromeroid type, parameres free distally. Larva - Hypopharyngeal sclerome subrectangular, beset with patches of setae and blunt, fleshy denticles. Ninth tergum expanded posteroventrally, forming the terminal urogomphal plate, bearing a pair of heavily sclerotized, 2- branched urogomphi, and two deeply pignented, heavily sclerotized urogomphal pits caudomesally, between the urogomphi; sternum nine bearing a single stout callosity on each anterolateral aspect. Remarls Fall (1915: 10) created the impression that Jacquelin-Duval (1859-63) was first to note apparent differences in abdominal structure between Pedilus and Corphyra. However, as Jacquelin—Duval (p. 364, footnote 1) clearly pointed out it was Lacordaire (1859: 37, footnote 2) who first commented on the apparent distinctions: En effet, M. Lacordaire dit avoir observe, chez trois exemplaires males, appartenant ‘a deux especes de l'Ame’rique du Nord, seules de ce pays lui e’tant connues, six arceaux ventraux ‘a l'abdomen, dont 1es deux derniers notablement plus grands que les autres. And, although Jacquelin-Duval stated, "MM. LeConte et Lacordaire ont rapporte au genre Pedilus un certain nombre d‘especes de l'Amerique du Nord, quui peut-Etre devraient appartenir ‘a un genre distinct (Corphyra Say)," neither 88 Lacordaire nor Jacquelin-Duval actually used the name Corphyra in a generic sense. It was, in fact, first elevated to generic status by LeConte (1867a). Abdullah (1969) erected a separate genus, N‘eopedilus, for p, parvicollis Fall. However, evidence presented in the discussion of phylogenetic relation- ships strongly supports the hypothesis that P. parvicollis and P. flabellatus are sister species and form a monophyletic species group. Furthermore, structural similarities between P. parvicollis and several Russian species of Pedilus support the hypothesis that the Palearctic faunal element is most closely related to the pumctuulatus-flabellatus lineage of the Nearctic element, to which p, parvicollis belongs. N eopedilus is therefore regarded as a synonym of Pedilus. The North American species of Pedilus are divided into six presumably monophyletic lineages or species groups based upon evidence presented above (see Intrageneric Phylogenetic Considerations). CHECKLIST OF NORTH AMERICAN PEDILUS SPEC- GROUPS AND SPECIB flabellatus group: 1. P. flabellatus (Horn) 2. P. parvicollis Fall mm m 3. P. abnormis (Horn) 4. P. dentatuus Abdullah 5. P. rusticus Abdullah 6. P. fenderi, sp. nov. 7. P. flexiventris Fall 8. P. cavatus Fall 9. P. jolmsonorum. sp. nov. 10. P. inconspicuus (Horn) 11. P. vittatus (Horn) 12. P. oregonus Fall 13. P. bardi (Horn) 14. P. punctulatus LeConte lewisi group: 15. P. picipennis Fall 16. P. elegans (Hentz) 17. P. granti, sp. nov. 18. P. lewisu’ (Horn) 89 90 labiatusgroup: 19- P.10ngilobusFall 20. P. crotchi (Horn) 21. P. labiatus (Say) 22. P. joanae sp. nov- 23- P. monticolus (Horn) numb-ism 24. P. lugubris(SaY) 25- P. cyanipennis Bland terminalisgroup: 26. P. canaliculatus (LeConte) 27- P. impressus (Say) 28. P. terminalis (Say) unplaced species: 29. P. serratus Fall 30. P. alticolus Fall ulcer-tee seas: Corphyra calypso Wickham ARTIFICIAL KEYS TO SPEC. OF NORTH AMERICAN PEDILUS A. Abdomen with six visible, freely articulated sternites; hind margin of last visible sternite (= 8th) usually emarginate; elytral organs (see p. 16C) usually present; param eres of genitalia usually extruded beyond apex Of mmen‘...OOOOOOOOOOOOOOOO0.0000000000000000000000000000000.0......OOOMALE (p91 ) A'. Abdomen usually with five visible, freely articulated sternites; hind margin MALES 1. l'. (1) 2. 2'. (2') 3. of last visible sternite (= 7th) usually entire, rounded; elytral organs absent; 8th abdominal sternite and genitalia usually retracted within abdomen. ..... . ............ ...FEMALES (p. 99 ) Antennae pectinate (Figure 23) to flabellate (Figures 18-19)..............2 Antennae strongly serrate (Figure 29), serrate (Figures 20-22, 25-26) or subfiliform (Figures 27-28)4 Antennae pectinate; pronotum ovate, widest at the middle; elytral apices slightly impressed; parameres each bearing a well developed, inwardly directed subapical tooth from the inner dorsal surface (Figure 87); distribution as indicated by Figure 161 crotchi (Horn) (p.183) Antennae flabellate; pronotum widest anteriorly (Figure 33); elytral apices not impressed; parameres lacking inwardly directed subapical tooth.3 Antennae flabellate beyond 2nd article (Figure 19); fused portion of parameres prolonged distally, forming acutely rounded medial 91 (l') (4) (5) 3'. 4'. 6'. 92 lobe between parameres (Figure 69); distribution as indicated by Figure l43parvlcollis Fall (p110) Antennae flabellate beyond 3rd article (Figure 18); parameres not separated by medial lobe (Figure 68); distribution as indicated by Figure 142flabellatus (Horn) (p105) Antennae weakly serrate, subfiliform (Figures 27—28); east of Rocky Mountains.5 Antennae serrate (Figures 20-22, 25-26, 29); mostly from Rocky Mountains to West Coast [exceptionsz joanae (Figure 163), labiatus (Figure 162)] 10 Elytra concolorous, iridescent purple or piceous to black...................6 Elytra piceous to black with creamy light yellow to testaceous apicess Elytra iridescent indigo blue-green to purple, each bearing a deep subapical and juxtasutural reniform impression; legs with coxae and femora piceous to black, remainder rufotestaceous; param eres conspicuously flattened dorsoventrally and tapered to a point distally (Figure 92); Appalachian Mountains from Virginia to southern Quebec (Figure 166)...............cymipennis Bland (p.214) Elytra piceous to black, impressed areas neither deep nor reniform; legs usually piceous to black, infrequently testaceous or black with lighter areas but not as above; parameres flattened or not, if conspicuously flattened, then abruptly narrowed distally (Figure 91); Widely fistributed mt Of kay Momtaimoooooeooooooooeoooeeooooo7 (6') (5') (8') 7'. 9'. 93 Elytral apices each bearing a preapical and juxtasutural ovate impression; elytral suurface coarsely and moderately sparsely punctate; parameres each bearing a single acute, dorsoanteriorly directed tooth-like process from the inner ventral suurface (Figure 94)impressus (Say) (p226) Elytral impression apical; elytral surface moderately coarsely and densely punctate to rugose; parameres (Figuure 91) lacking tooth- like “meaOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOlWis (say) (p.206) Head and pronotum dorsally microsculptured (Figure 34), surface appearing isodiam etrically patterned; pronotum bearing longitudinal sulcus mesally; parameres lacking prominent tooth— like processes; distribution as indicated in Figure 167.000.000.00000000 00.00.00.000....OOOOOOOOOO. cmalicmaw (Leconte) (p218) Dorsal surface of head and pronotum finely and sparsely punctulate; pronotum not sulcate; parameres each bearing a single prom inent, subapical tmttl-like mm (Fines 83, 95)....OOOOOOOOOOOOOOOOOOOOOOOOOOO9 Abdominal sternites 3-6 piceous to black, 7-8 testaceous to rufotestaceous; elytra moderately coarsely, densely punctate; apices of parameres excurved; each paramere bearing a prominent dorsomesally projecting tooth—er structure from the inner ventral surface (Figure 83); distribution as indicated by Figure . l57000OOOOOOOOOOOOOCO...0.0.0.0000...OOOOOOOOOOOO0.000000000 elegm (HentZ) (p.165) Abdomen usually concolorous black, distal portions of sternites 7-8 occasionally testaceous to rufotestaceous; elytra coarsely, moderately sparsely punctate; parameres straight, apices bluntly pointed; each paramere bearing a prominent anteromesally (4') 10. 10'. (10') 11. 11'. (11) 12. 12'. (12) 13. 13'. (13) 14. 14'. 94 directed tooth-like process from the inner surface (Figure 95); distribution as indicated by Figure 169 ..terminalis (Say) (p.231) Tooth of tarsal claw subequal in length to claw itself, truncate distally (Figure 47); distribution as indicated by Figure 160 ..longflobus Fall (p.180) Tooth of tarsal claw small (Figure 46) or, if large (Figures 42-45), acutely pointed distally and never more than 1/2 - 2/3 length of claw itselfll Tooth of tarsal claw small (Figure 46)12 Tooth of tarsal claw well developed (Figures 42-45).........................22 Abdominal sternite 7 strongly deflexed ventrally.............................l3 Abdominal sternite 7 norma1.l6 Elytral apices weakly, juxtasuturally impressed or not modified at all; elytra concolorous rufopiceous to black or black with narrow testaceous sutuural margin distallyl4 Elytral apices abruptly. deeply, obliquely impressed; elytra bicolorous, black with yellowish to stramineous apices or stramineous to testaceous with impressed areas, sutural margins and lateral Frontoclypeal surface of head finely, densely punctate; parameres narrow, acutely rounded apically (Figure 73); distribution as indicated by Figure l47..............................fenderi sp. nov. (p124) Frontoclypeal surface of head coarsely, densely punctate; parameres wide, obliquely truncate apically (Figure 72); distribution as indicated by Figure l46...........................rusticus Abdullah (p.121) (13') 15. 15'. (12') 16. 16'. (16) 17. 17'. (16') 18. 18'. 95 Elytra stramineous to testaceous with impressed areas, sutural margins and lateral margins black; distribution as indicated by Figure l49cavatus Fall (p.130 ) Elytra black with yellowish to stramineous apices; distribution as indicated by Figure l48.............................flexiventris Fall (p127) Median lobe with sides abruptly convergent apically, apex produced into acutely pointed mesal tooth or spine-like process (Figures 107, 109); elytral apices not modufued.l7 Median lobe with sides gradually tapered distally (Figures 111-114, 116); elytral apices modified or not18 Apical process of median lobe short, stout, tooth-like (Figure 109); parameres each bearing an inwardly projecting, serrulate, molar- like process (Figures 77, 110); distribution as indicated by Figure lslinconspicuuus (Horn) (9138) Apical process of median lobe elongate, spine-like (Figure 107); parameres each bearing a well developed dorsoanteriorly directed subapical tooth from the inner ventral suurface (Figures 76, 108): distribution as indicated by Figure 150 johnsonorum, sp. nov. (p.133) Parameres excurved subapically, each forming a well developed, outwardly projecting tooth and bearing a broad inwardly projecting tooth from inner margin (Figure 82); elytral apices not modified; distribution as indicated by Figure 156 pucipennus Fall (p.162) Parameres not excurvate, bearing only an inwardly projecting subapical tooth (Figures 78-81); elytral apices modified or notOOOOOOOO......OOOOOOOOOOOOOOOOOOOOOO0...... ......... O ...... O OOOOOOOOOOOOOOOOOOOOOOOOO 00.19 96 (18') 19. Elytra caudate and upwardly curved apically; apex of elytra impunctate, somewhat swollen; distribution as indicated by Figure 154....... ........... .. ........... ...... .. ....... .............bardi (Horn) (p352) 19'. Elytra neither caudate nor upcurved apically; apex of elytra impunctate and swollen or not20 (19') 20. Elytra widely flattened along distolateral margins; distribution as indicated by Figure 153 ....... ..... . ...... oregonus Fall (p.149) 20'. Elytra not broadly impressed along distolateral marginle (20') 21. Elytral apices impunctate, somewhat swollen; subapical tooth of parameres acute (Figure 81); distribution as indicated by Figure l55............ ......... ........ . .............. punctulatus LeConte (p156) 21'. Elytral apices not modified, neither impunctate nor swollen; subapical tooth of parameres stout, obtuse (Figure 78); distribution as indicated by Figure 152 ...... . ...... . ................. vittatus (Horn) (p.145) (11') 22. Elytral apices conspicuously impressed and/or swollen, the modified area impunctate and shunmg24 22'. Elytral apices neither impressed nor swollen, at most flattened along sutural angles, the flattened region not impunctate and shunmg23 (22') 23. Metathoracic tibiae slightly arcuate, inner tibial surface flattened with posterior margin obliquely carinate, thus giving tibiae slightly twisted appearance; parameres bluntly rounded and slightly convergent distally, each bearing a small subapical tooth from the inner margin (Figure 70); widely distributed west of the Rocky Mountains (Figure 144)...... .............. abnormis (Horn) (p114 97 23'. Metathoracic tibiae normal, neither arcuate nor twisted; parameres acutely rounded distally, each bearing a well developed subapical tooth from the inner margin (Figure 71); distribution as indicated by Figure 145 dentatus Abdullah (p.119) (22) 24. Parameres excurved subapically, each forming an outwardly projecting process; inner aspect of each paramere bearing an acute preapical tooth (Figures 84—85); elytra impressed preapically, swollen surrounding impressed area, impressed and swollen areas impunctate, shunmg25 24'. Parameres with apices variously tapered, never excurvate or bearing outwardly projecting process; if inner aspect of each paramere bears a tooth, it is acute and subapical (Figure 96) or poorly defined and obtuse (Figures 88-90); modification of elytral apices varuab1e26 (24) 25. Elytra rufotestaceous with apices black and preapical yellowish-orange reniform spot; parameres each bearing a single, small, dorsoanteriorly projecting spine from the ventrointerior surface (Figure 84); known only from King's Canyon National Park, CA (Figure 158) granti,sp, nov. (p. 171) 25'. Elytra black with swollen preapical region black or yellowish, or elytra entirely testaceous; parameres each bearing a single, well developed, anteromesally curved subapical tooth from inner margin (Figure 85); New Mexico and eastern Arizona to southeastern Wyoming (Figure 159).. ...... . ..... . lewisi (Horn) (p.174) 98 (24') 26. Elytral apices each with a single, well developed juxtasutural impression which is slightly more than twice as long as its maximum width; parameres each bearing a preapical tuft of anteromesally directed spinulae along the inner margin (Figure 97); distribution as indicated by Figure 171 ....alticolus Fall (p.240) 26'. Modification of elytral apices variable, impressed area never elongate and narrow; inner aspect of each paramere bearing an acute subapical tooth (Figure 96) or a poorly defined, obtuse prominence (Figures 88-90)27 (26') 27 . Elytra weakly, flatly impressed apically, slightly swollen anterad of impressed area; inner aspect of each paramere bearing an acute subapical tooth (Figure 96); distribution as indicated by Figure 170 serratus Fall (92%) 27'. Elytral apices each with a subcircular impression, elytra slightly swollen surrounding impressed region; inner aspect of each paramere bearing a poorly developed, obtuse preapical prominence (Figures 88-90)28 (27') 28. Elytra with sutural angle prolonged into a dentiform process (Figure 48); eastern North America from East Coast to Montana, Florida to southern Michigan (Figure 162) labiatus (Say) (p.188) 28'. Elytra with apices acutely angulate but sutural angle not produced to form dentiform process; western North America from West Coast to MinnemtaOOOOOOOO..OOOOOOOOOOOOOOOOOOOOOOOOOOOOO0.000000000000000.00000000000000000zg 99 (28') 29. Parameres with ventral sinus usually well developed (Figure 89); distribution as indicated by Figure 163........joanae, sp. nov. (p. 94) 29'. Parameres totally lacking ventral sinus (Figure 90); distribution as indicated by Figure l64..........................monticolus (Horn) (p.200 FEMALES (Region 1, see Figure 9) [Note: females of dentatus, fenderi and (l) 2. (2) 3. 3'. (2') 4. 4'. (4) 5. 5'. grant! have yet to be discovered.) Tooth of tarsal claw small (Figure 46)2 Tooth of tarsal claw well developed (Figures 42-43, 45)....................ll Pronotum widest anteriorly (Figure 33)3 Pronotum ovate, widest at muddle4 Dorsal surface of head moderately finely, sparsely pumctate; distribution as indicated by Figure 142......flabellatus (Horn) (p.115) Dorsal surface of head coarsely, moderately densely punctate; distribution as indicated by Figure l43.........parvicollis Fall (p. 110) Entire dorsal surface of head and pronotum very coarsely densely punctate............. ........ 5 Dorsal surface of head and pronotum finely and sparsely to moderately coarsely and densely punctate, or head punctation variable with coarse, dense punctation limited to frontoclypeal and postocular aspects of head6 Lateral extension of ventral bacculus rod-like (Figure 63) .bardi (Horn) (p. 152), oregonus Fall (p. 149), punctulatus LeConte (p. 156 Lateral extension of ventral bacculus reduced, visible only as lightly sclerotized baccular plate (Figure 64)..........vittatus (Horn) (p.145) (4') (6') (7') (8') (9') 6'. 7'. 8'. 9'. 10. 100 Dorsal surface of head densely covered with elongate yellowish-golden setae; elytra stramineous to testaceous, apices, sutural margins and lateral margins black; distribution as indicated by Figure l49cavatus Fall (p.130) Head sparsely covered with short to moderately elongate setae and elytra testaceous to black, or head moderately densely covered with yellowish setae and elytra black.7 Hind margin of 7th abdominal sternite conspicuously, acutely produced (Figure 57); distribution as indicated by Figure 146 Hind margin of 7th abdominal sternite broadly (Figures 59-60) to narrowly (Figures 58, 61) rounded, not acutely produced..............8 Head and scutellum orange to reddish-orange or orange suffused with black; distribution as indicated by Figure 150 jolmsonorum, sp. nov. (p.133) Head and scutellum testaceous to b1ack.9 Postocular surface of head very coarsely, densely punctate; distribution as indicated by Figure 151 inconspucuus (Horn) (p.138) Postocular surface of head finely, sparsely punctate.......................10 Surface of head covered with short setae; elytra testaceous to black; pronotum rufopiceous to black (rarely orange); distribution as indicated by Fime l56ooooooeeoocoosooooooeoooeooooepiCipeMis Fm (p 162) 10'. Surface of head covered with moderately elongate, yellowish setae; elytra black; pronotum yellowish-orange to orange; distribution as indicated by Figure l48.............................flexiventris Fall (p.127) 101 (1') 11. Elytral apices slightly, acutely produced (Figure 49); distribution as indicated by Figure 171...............,..,..,..,........alticolus Fall (p. 700) ll'. Elytral apices rounded, not at all produced.12 (11') 12. Dorsal surface of head finely and moderately sparsely punctate........l3 12'. Dorsal surface of head moderately coarsely and densely punctatel4 (12) 13. Antennal articles 1-2 testaceous to rufotestaceous; distribution as indicated by Figure 160 ..... longilobus Fall (p. m) 13'. Antennal articles 1-2 black; widely distributed west of the Rocky Mountains (Figure 14001110171113 (Horn) (p.114) (12') 14. Antennae serrate to strongly serrate (Figures 24, 30)l5 14'. Antennae weakly serrate, subfiliform (Figure 26) Joanne, sp. nov. (p.194), monticolus (Horn) (1)-20°) (14) 15. Femora and venter of abdomen concolorous rufopiceous to black; distribution as indicated by Figure 17o...........serratus Fall (p4236 ) 15'. Femora and venter of abdomen variegated, testaceous to amber and piceous to black; distribution as indicated by Figure 161 ..... . .......... crotchi (Horn) (p.183) FEMALES (Region 2, see Figure 9) l. Tooth of tarsal claw small (Figure 46)2 l'. Tooth of tarsal claw well developed (Figures 42-43, 45)4 (1) 2. Dorsal surface of head and pronotum moderately coarsely, sparsely to moderately densely punctate; distribution as indicated by Figure l56.00.0000...OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO. picipemm Fall (p.162) 102 2'. Dorsal surface of head and pronotum very coarsely, densely punctate3 (2') 3. Elytral apex acute (Figure 50); distribution as indicated by Figure ' 154..bardu (Horn) (p.152) 3'. Elytral apex obtusely rounded (Figure 51); distribution as indicated by Figure lszvuttatus (Horn) (p.145) (1') 4. Elytra with sutural angle prolonged into a dentiform process (Figure 48); eastern North America, East Coast to Montana, Florida to southern Michigan (Figure l62)......................labiatus (Say) (p.188) 4'. Elytra with sutural angle rounded, not produced.5 (4') 5. Dorsal surface of head and pronotum moderately coarsely and densely puunctate; elytra testaceous to black; distribution as indicated by Figure 163 joanae, sp. nov. (p.194) 5'. Dorsal surface of head and pronotum finely and moderately sparsely punctulate; elytra piceousto black.6 (5') 6. Antennae entirely piceous to black; widely distributed from the Rocky Mountains to the West Coast (Figure 144)....abnormis (Horn) (p.114 6'. Antennae rufopiceous to black with distal portions of articles 1-2 usually testaceous; distribution as indicated by Figure l59.00.000.00...0.000000000000000000000000000000000000000000000000 lewui (Horn) (p. D4) FEMALE (Region 3, see Figure 9) l. Elytra iridescent indigo blue-green to purple; legs 'with coxae and femora piceous to black, remainder rufotestaceous; Appalachian Mountains from, Virginia to Southern Quebec (Figure 166)OOOOOOOOOOOOO0.0000000000000000.00.000.000...0......”wipemw Bland (p. 214) (1') (2') (3') (4') (5') l'. 2'. 3'. 4'. 5'. 6'. 103 Elytra testaceous to black; legs usually concolorous, testaceous or piceous to black, infrequently black with lighter areas but not as above; widely dustrubuted.2 Head and pronotum dorsally microsculptured (Figure 34), surface appearing isodiametrically patterned; pronotum bearing longitudinal sulcus mesally; distribution as indicated by Figure 167 canaliculatus (LeConte) (p. 213 Dorsal surface of head and pronotum finely and sparsely to moderately coarsely and densely punctate; pronotum not sulcate; widely Elytra with sutural angle prolonged into a dentiform process (Figure 48); eastern North America, East Coast to Montana, Florida to southern Michigan (Figure l62).....................labiatus (Say) (p. 133) Elytra with sutural angle rounded, not produced.4 Dorsal surface of head and pronotum moderately coarsely and densely punctate; elytra testaceous to black; distribution as indicated by Figure 163 joanae, sp. nov. (p.194) Dorsal surface of head and pronotum finely and sparsely punctulate; elytra piceous to black5 Elytra coarsely, moderately densely punctate; pronotum usually transversely ovate umpressus (Say) (p. 226, terminalis (Say) (p. 231) Elytra moderately coarsely, densely punctate; pronotum ovate...........6 Lateral extension of ventral bacculus rod-er (Figure 63) lugubris (Say) (p.206 Lateral extension of ventral bacculus reduced, visible only as lightly sclerotized baccular plate (Figure 64)... ..... .elegans (Hentz) (p. 165) THE FLABELLATUS GROUP Diagnosis Males of this group are characterized by their flabellate antennae (Figures 18-19). In both the males and females, the eyes are coarsely faceted and the pronotum is distinctly wider anterad of the middle. Remarks On the basis of anatomical characters and what little may be inferred regarding bionomics, the two species comprising this group depart greatly from all other Pedilus. Although the antennal modificatious of male serratus and crotchi may suggest a transformation series, characters associated with the genitalia, and structure of the eyes, pronotum and tarsal claws fail to support such a hypothesis. Thus, the antennal apomorphies must be accounted for though evolutionary convergence. The flabellatus group has a western montane distribution pattern. 104 105 l. Pedilus flabellatus (Horn) (Figures 18, 33, 68, 98, 142) Corphyra flabellate Horn, 1883: 306-307. Type locality: "Western Nevada." Lectotype, designated by Abduullah, 1962: 219, 0' , #3036, MCZC. . Pedilus (Corphyra) flabellatus Horn; Pic, 1911: 12 (catalog). Pedilus flabellatus Horn; Fall, 1915: 15, Figures 1, 6c, 7 (distribution). Pedilus flabellatus (Horn); Leng, 1920: 161 (catalog); Abdullah, 1962: 219 (designation of lectotype); Abduullah, l964d: 19-20, Figures 19-24, map 3 (distribution, bionomics). Type Information In the original description, Horn stated that he had seen six specimens, all from western Nevada, one of which was a male with "pale" elytra. Five specimens were examined from the Horn collection (2 0'0' , 3 99 ); one of the males is somewhat teneral and may be the specimen with "pale" elytra. One specimen was also discovered in the LeConte collection, bearing the same "Nev" label, presumably the sixth specimen of the syntype series. Thus, the lectotype designated by Abdullah should be valid. He should, however, have labelled the other five syntypes with the designation "paralectotype" as recommended by the International Code of Zoological Nomenclature (1961-1964: 81). Abdullah (1964d: 20) incorrectly referred to the lectotype as a holotype. 106 Doa'iption Male: Length 4.5-7 mm. Color of prothorax and legs testaceous to orange; head, remaining exposed regions of thorax including elytra, and visible abdominal sternites ruufopiceous to black. Head sparsely setose, finely and sparsely punctate; antennae (Figure 18) flabellate beyond 3rd article, eyes large, rather coarsely faceted. Pronotum (Figure 33) widest anteriorly, its surface sparsely setose, finely and sparsely punctate. Legs with prothoracic and mesothoracic tarsi no more dilated and tomentose beneath than those of metathoracic legs; tarsal claws each with a small, poorly developed basal tooth. Elytral suurface finely and moderately sparsely punctate, lacking pruuinose luster; elytral apices not modified. Posterior margins of abdominal sternites 7-8 nearly straight; 8th abdominal segnent usually retracted within 7th. Parameres (Figure 68) fused proximally, widely separated and subparallel along distal 1/4; median lobe (Figure 98) excavated ventrally, apex narrowly rounded. Female: As for male except antennae serrate. Lateral extersion of each ventral bacculus well developed. Distribution (Figure 142) PACIFIC NORTHWESTtBRITISH COLUMBIA : Trinity Val.; 25-V-1928; CASC(1d'). Trinity Val.; 12-V11-1928;CASC, CNCI (2 o‘o' ). Annis Bay, Nelson Island; 3l-VII-1927; CASC (l d' ). OREGON : 7 mi. N.W. Roseburg, Douglas 00.; 10-VII-l964; JSCC (l 9 ). Bridge Camp, Myrtlewood; 27-VII/4-VIII-1955; BMUW, ODAC (l o' , 2 99 ). Rogue Rv. N.F., Beaver Sulphur F.C.; 10-VIlI-l950; BMUW (l 9 ). Homestead Inn, Mt. Hood; 2-VlI-l927; CASC (l d' ). Deschutes 00., SW 107 Sec. 15, T168, R93, 10 mi.SSE Sisters; 19-VII-1978; ODAC (10' ). Grants Pass; 2- VII-l967; ODAC (19). Wash. Co., 6 mi. NE Newberg; 2-VIlI-1968; ODAC (1 9 ). Allen Springs, Metalius Riv., Willamette N.F.; 10-Vll-1969; DYCC (l o' ). Murphy Crk, 2 mi. 8. Murphy, Josephine Co.; 3-Vll-l969; DYCC (l 0' ). CALIFORNIAN: CALIFORNIA: Carrville, Trinity Co.; s-vn-lsso; CASC (19). Plumas Co., Mohawk; ll-VlI-l957; CASC (ld'). Plumas Co., Mohawk; 12-VlI-l957; CASC (10‘). Chester; ll-VII-1951; OSUC (4 0' 0' ). Idyllwild, San Jacinto Mts.; 20-VI-1940; CASC (10' ). Clio, Plumas Co.; 14-VI-1914; CASC (l d ). Huntington Lake, Fresno Co., 7000 ft.; 8-VII-1919; CASC (1 0' ). Huntington Lake, 19-V11-1936; UCDC (1d ). Facht, Lesson Co.; 20-VI-1921; CASC (1 0' ). Tallac; "July"; CASC (l o' ). Fallen Leaf Lake, L. Tahoe; 20-v1-1915; CASC (1 0'). San Jacinto Mts, Marion Camp; 6-Vll-l950; UCDC (1 9 ). Hanna Flats, San Bernardino Co.; 20- VII-1952; LACM (19). "Lee Vinin 5", Mono Co.; 4-VII-l962; CDAE (19 ). McCloud; 29-v1-1914; CASC (l o' ). Hat Creek P.O., Shasta Co.; 2041-1955; CISC (10‘). Hat Creek P.O., Shasta Co.; 21-V1-l955; CISC (10'). Cedar Pass Cpgd., Modoc Co., 5900'; 23-VII-1968; CISC (10' ). Sand Flat, Alt. 5500 ft.; 24- v1-1930; UCRC (l 9 ). Inyo Nat. For.; 12-v11-1957; MCZC (l 0'). Lake Tahoe; 17-21/VlI/1897; MCZC (l 0' ). "Sugar Pine"; [no date]; MCZC (l 0' ). Sisson; 24- V1-1898; MCZC (10' ). Cold Crk. F.C., Tahoe N.F., 5500'; 23-VI-1968; DYCC (1 d).Tannery Gulch F.C., Claire Engle Lake; 24-Vl-1967; DYCC (1 0' ). Humboldt Co., Richardson Grv., 7 mi. 8. Garbervle; l6-VlI-l977; DYCC (l 9 ). Trinity Co., Shasta-Trinity N.F., Tannery Gulch Cmpgd., 5000'; 21-22/VI/ 1978; DYCC (20‘). Sagehen Crk., Nevada Co.; 25-VI-1966; DCCC (1 d' ). NEVADA: [State label only]; MCZC (10' , 1 9 ). Specimens examined: 36 do', 13 99 . 108 Diagnosis The flabellate antennae and anteriorly widened pronotal configuration serve to distinguish males of flabellatus and parvicollis from all other Species of Pedilus. The males of crotchi bear some resemblance due to their pectinate antennal structure. However, in crotchi the pronotum is ovate and widest at the middle. In flabellatus, the antennae are flabellate beyond the third article; they are flabellate beyond the second article in parvicollis. Females of flabellatus and parvicollis differ from all other Pedilus by the following set of characters: basal tooth of tarsal claw small; pronotum widest anteriorly. The dorsal surface of the head is moderately finely, sparsely punctate in flabellatus females and coarsely, moderately densely punctate in those of parvicollis. Remarks The specimens examined are relatively homogeneous. The most notable source of intraspecific variation is associated with the elytra, which vary in coloration from rufopiceous to black; 2-3 specimens with nearly testaceous elytra appear to be somewhat teneral. Bionounics Abdullah (l964d) noted that this species had been collected from Pinus monticola; three males were examined from Trinity Valley, British Columbia which bear this label. Both males and females have been collected from lights at night (Oregon and California records), indicating that flabellatus may be crepuscularly or nocturnally active. The morphology of the eyes and antennae, especially in the 109 male, further supports this notion. James and Janice Johnson collected a Single female at "black light" in the Richardson Grove redwood stand of Humboldt County, California. The Site was located in a patch of Arbutus and U mbellulario along the Eel River; no specimens were observed at cantharidin bait during the day. James Johnson and Joan Young each collected single males in flight at Tannery Gulch campground in the Shasta-Trinity National Forest. Both Specimens were taken between 9:30-10:30 a.m., thus supporting a crepuscular rather than strictly nocturnal activity hypothesis. Once again, no Specimens were observed at cantharidin bait during the day. 110 2. Pedilus pavicollis Fall (Figures 19, 33. 69, 99, 143) Pedilus parvicollis Fall, 1919: 216; Long and Mutchler, 1927: 29 (catalog); Fall, 1929: 13-14 (generic position); Abdullah, 1962: 219; Abdullah, l964d: 20-21, Figures 25—30, map 4. Type locality: "Monache Meadows, Tulare Co., [California] 9,000 to 8,300 feet." Holotype, 0‘ , #24321, MCZC. Dendroides parvicollis of Van Dyke, 1928: 260 (generic position). Neopedilus parvicollis of Abdullah, 1969: 359 (generic position). Desaiption Male: Length 6.5-9 mm. Color of mouthparts, labrum, anterior portion of clypeus, prothorax, prothoracic coxae, and scutellum testaceous to reddish- orange, remainder of head, exposed regions of thorax including legs and elytra, and visible abdominal sternites rufotestaceous to black. Head sparsely setose, coarsely but sparsely punctate; antennae (Figure 19) flabellate beyond 2nd article; eyes large, coarsely faceted. Pronotum (Figure 33) widest anteriorly, its surface sparsely setose, moderately coarsely but sparsely punctate. Legs with prothoracic and mesothoracic tarsi no more dilated and tomentose beneath than those of metathoracic legs; tarsal claws each with a small, poorly developed basal tooth. Elytral surface moderately finely and densely punctate to rugose, lacking pruinose luster; elytral apices not modified. Posterior margin of 8th abdominal sternite subtruncate. Parameres (Figure 69) fused proximally, widely separated and subparallel along distal 1/2, fused region produced distally as an acutely rounded medial process; median lobe (Figure 99) slightly excavated ventrally, apex narrowly rounded. 111 Female: As for male except antennae serrate. Lateral extension of each ventral bacculus well developed. Distribution: (Figure 143) CALIFORNIAN: CALIFORNIA: Sequoia Park; VII-1906; LACM (1 0‘ ). Roumd M'd'w, Giant Forest; "June"; CNCI (l 0' ). Rouund M'd'w, Giant Forest; "July"; CASC (l d ). Young Lake, Tuol. Co.; 19-VllI-l947; CISC (1 9 ). Gray de., Lot 13, Tulare Co.; 3-VIl-l9l3; CASC (1d). Seafford de, Tulare Co.; 11- VlI-l910; CNCI (10' ). Mt. Mitchell; VIII-1914; CASC (l o‘ ). Fresno Co., Rea Lake, Alt. 10,500 ft.; 20-v11-1910; CASC (1 o' ). Fresno Co., Bullfrog L., Alt. 10,600 ft.; 20-26/VIII/ 1908; CASC (1 0' ). Huckleberry Meadow, 6,500 ft., "H- 1050 Dendroides sierrae (ms) Van Dyke"; l-Vlll-l909; CASC (1 o' ). Ediza Lake, Mono Co.; 5-16/VllI/1958; LACM (2 0'0' ). Tulare Co., Monache Meadows, 8-8300 ft.; 15-VlI/l3-Vlll; MCZC (1 0' ). Tulare Co., Monache Meadows, 8-8300 ft.; 28- VI/l3-VIll; MCZC (1 o’ ). lnyo Co., Poling; "Aug, 1922"; MCZC (19). Round M'd'w, Giant Forest; "June"; MCZC (l 0' ). Kings Canyon Nat'l Park, Acc. No. 361; 3l-V11-1952; SJSC, DYCC (3 o'o' ). [no state label, presumably California], Lot 908, Sub 1457; [no date]; CUIC (2o). Specimens examined: 19db‘, 2929. Diagnosis The males of parvicollis are unique in having the antennae flabellate beyond the second article. This condition is approached by males of flabellatus, in which the antennae are flabellate beyond the third article, and males of crotchi which have delicately pectinate antennae. Females of parvu’collis and flabellatus are easily distinguished by the anteriorly widened pronotal configuration; the pronotum of other species is ovate and widest at the middle. 112 In female parvicollis, the dorsal surface of the head is coarsely and moderately densely punctate; it is moderately finely, sparsely punctate in flabellatus females. Remarks The small number of specimens currently known are quite homogeneous in terms of size, color and structure, the primary difference being in the sexual dimorphism of the antennae. In several respects, most notably the structure of the antennae and eyes, this species strongly resembles the pyrochroine genus, Dendroides. In fact, these similarities led Van Dyke (1928) to conclude that parvicollis should be transferred to Dendroides. Although his contention was soundly rejected by Fall (1929), the fact remains that the two share some interesting characters. In Dendroides, a nocturnally active group, the development of the antennae and eyes is thought to represent an evolutionary response to functioning (e.g. mate location) in low light. Nothing is known about the bionomics of pm'vu'collis, but the above characters suggest that parvicollis may also be a nocturnally or crepuscularly active species, its Similarity to Dendroides being accounted for by evolutionary convergence of sensory mechanisms. Supportive evidence comes from P. flabellatus, the probable sister Species of parvicollis. lt possesses similar modifications of the antennae and eyes, and has been taken on several occasions at lights. Bionomics Virtually nothing is known about the natural history of parvicollis (but see remarks, above). THE PUNCTULATUS GROUP Diagnosis The members of this group are defined by the presence of paired serrate structures which are associated with the posteroventral surface of the median lobe (Figures 100-107, 109, 111-114). Remarks Within this presumably monophyletic lineage two distinct assemblages of species were identified: the cavatus subgroup is comprised of P- rusticus, P- fenderi, P. flexiventris and P. cavatus while the punctulatus subgroup includes P. vittatus, P. oregonus, P. bardi and P. punctulatus, This is a western group with five species endemic to California and not a single species found east of the Rocky Mountains. 113 ‘1 114 3. Pedilus calla-unis (Horn) (Figures 2, 70, 100, 144) Corphyra abnormis Horn, 1874: 40; Horn, 1883: 307, pl. 9, Figure 17 (distribution). 'Iype locality: "California." Holotype, 0' , #3032,MCZC. Pedilus (Corphyra) abnormis Horn; Pic, 1911: ll (catalog). Pedilus abnormis Horn; Fall, 1915: 16-17, Figures 5, 6b, 8a, 8b (description of female; distribution). Pedilus infectus Fall, 1915: 17-18, Figures 6b, 8a, 8b; Abdullah, 1962: 219; Abdullah, 1966b: 187-188, Figures 56-61 (distribution). Type locality: "San Jacinto Mts. in Southern California." Holotype, o' , #24317, MCZC. NEW SYNONYM. Pedilus abnormis (Horn); Lang, 1920: 161 (catalog): Abdullah, 1962: 219; Abdullah, 1966b: 182-187, Figures 48-55 (distribution, bionomics). Description Male: Length 5-8 mm. Color of head, ventrally exposed portions of mesothorax and metathorax, legs, elytra and abdominal sternites 3-8 piceous to black; pronotum usually concolorous red-orange or piceous to black, less frequently red—orange suffused with black; prosternum usually piceous to black with red—orange anterior margin, less frequently concolorous piceous to black or red-orange. Head covered with moderately elongate erect setae, postocular aspect of genae finely and sparsely to densely puunctate. Antennae serrate, articles 1-2 frequently more darkly pigmented and shining than remaining segments. 115 Pronotum ovate, covered with moderately elongate, retrorsely semierect setae, suurface finely and sparsely punctate. Legs with metathoracic tibiae Slightly arcuate, inner tibial surface flattened and bearing an obliquely carinate posterior margin, thus giving the tibiae a slightly twisted appearance; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws strongly toothed at base. Elytral suurface densely punctate to rugose, lacking pruinose luster; elytral apices not modified. Abdomen with 6th sternite produced posteromesally, thus forming a broad lobe, the distal margin of which is weakly bisinuate to nearly straight; 7th sternite tapered posteriorly, broadly and moderately to shallowly emarginate along distal margin; 8th sternite emarginate along hind margin. Parameres (Figure 70) somewhat dorsoventrally flattened, fused proximally, free and subparallel along distal 1/3. each bluntly rounded and very Slightly convergent apically, each bearing a single small inwardly projecting subapical tooth from the inner ventral surface; median lobe (Figure 100) bearing paired, distally free and movable serrated appendages ventrally, apex tapering to a blunt point. Female: Differs from male by lacking specializations of metathoracic tibiae; tarsi of prothoracic and mesothoracic legs scarcely more tomentose beneath than those of metathoracic legs; elytral surface pruuinose or not. Abdominal sternites 6-7 normal, 8th sternite tapered with hind margin entire; lateral extension of each ventral bacculus well developed. Distribution (Figure 144) PRAIRIE: AB. MT; MOUNTAIN: co, UT, WY; PACIFIC NORTHWEST: ID, on; CALIFORNIAN: CA, NV. [AMNH, BMNH, BMUW, CASC, CDAE, cusc, CNCl, CUIC, DEFW, DYCC, DZEC, EMUS, FMNH, GHNC, JBJC, JSCC, LACM, 116 MCZC, MSUC, NHMB, ODAC, OSUC, OSUO, PMNH, PURC, SEMC, UASM, UCDC, UCEC, UCRC, UMMZ, UMRM, USNM, WSUCJ Specimens examined: 1259 60' , 262 99 . The majority of collections have been made between 20 June and 10 July. Diagnosis Males of abnormis may be easily distinguished from all others in the genus by the uunique structure of the metathoracic tibiae, the shape of the sixth abdominal sternite, or the structure of the parameres (Figure 70). The females share the following set of characteristics with those of lewisi and longilobus: dorsal surface of head and pronotum finely and moderately sparsely punctate; basal tooth of tarsal claw well developed; elytral apices rounded, not acutely produced posterad. In abnormis, the antennae are entirely piceous to black, whereas articles 1-2 are testaceous to rufotestaceous and the remaining segnents piceous to black in longflobus. Frequently females of lewisu' have the distal portions of antennal segnents 1-2 testaceous while the remainder of the antennae are rufopiceous to black. Remarks lntraspecific color variation is associated primarily with the prothorax. While a red-orange pronotum is by far the most common condition throughout the range of this species, pronotal color in a number of Specimens varies from red-orange tinted with piceous to black pigmentation to entirely black. The majority of Specimens possessing a black pronotum come from the Rocky Mountain region, but they are not limited to the region and there is no evidence to suggest an East-West cline. 117 Although no attempt has been made to analyze prothoracic color variation in terms of other possible clines, the distribution of darker specimens supports the speculation that elevation may be a significant factor, with darker pigmentation correlated to higher altitudes. I have examined the holotype of Fall's infectus along with 21 additional specimens from the San Jacinto Mountains and can find no consistent characters to separate these from abnormis. Most of the characters Fall used to distinguish infectus from abnormis relate to the small size (5.5-5.75 mm.) of the single pair he based the description on; his series of abnormis ranged from 6-8 mm. The series of abnormis examined ranges from 5-8 mm. and in the male, the degree of antennal serration, tarsal dilation, and metathoracic tibial modification is correlated rather closely to Size. Fall also noted that his female of infectus, "shows no trace of the pruuinose lustre so characteristic of typical abnormis." This character, too, is variable. Females from Colorado and Wyoming usually lack elytral pruinosity and several have been seen from Oregon and California (Inyo and Tuolumne counties) which show no trace of a pruinose luster. Bionomics Abdullah (1966b) noted that one specimen had been collected from a Sphagnum bog, while another was collected on Ceanothus integerrimus. Label data indicate that specimens from Alpine County, California, were taken on Salix and swept from grass; specimens were collected from Ceanothus in El Dorado County and from Pinus jeffreyi and "low weeds" in Riverside County, both in California. Numerous collections were made in Nevada County, California, from malaise traps placed in meadows. 118 Males of this species have been observed and collected in abuundance at cantharidin in Albany County, Wyoming, (Figure 2) and numerous locations in the Sierra Moumtains of California. The author and his wife collected numerous specimens of abnormis at and near cantharidin bait in Mariposa County; peak activity was noted between 4:00-4:30 p.m. Montane meadows and stream margins proved to be the most productive locations for the baits in both California and Wyoming (personal observation; James Johnson, in litt.). The ant, Formica lasiodes Emery, was observed dragging abnormis males from the vicinity of cantharidin bait placed at the margin of an alpine meadow at Yuba Pass, Sierra County, California (James Johnson, in litt.; ant determined by Duane Flynn). 119 4. Pedilus mm Abdullah (Figures 71, 101, 14s) Pedilus dentatus Abdullah, l964b: 163, tb. 7, Figures 71-74. Type locality: "Carrville, Trinity County, California." [Holotype, of , not examined, apparently lostJ Type Information Abdullah listed the California Academy of Sciences as repository for the holotype. Dr. Kavanaugh has informed me that he has been unable to find either the specimen or any record that it was ever returned to the California Academy of Sciences. The single paratype, o' , "Brit. Mus. 1964-136," BMN H, was examined. Description Male: Length 7.5-8 mm. All visible sclerotized areas rufopiceous to black. Head covered with moderately elongate semierect to erect setae, postocular aspect of genae finely and sparsely punctate; antennae serrate. Pronotum ovate, covered with moderately elongate semierect setae, surface finely and sparsely punctate, bearing a short longitudinal canaliculus mesally. Legs with prothoracic and mesothoracic tarsi conspicuously tomentose beneath: tarsal claws strongly toothed at base. Elytral surface densely punctate to rugose, lacking pruinose luster; apex of elytra flattened along sutural angle. Hind margin of 6th sternite truncate, that of 7th sternite subtruncate, 8th sternite with hind margin slightly emarginate mesally. Parameres (Figure 71) fused proximally, free and closely subparallel along distal 3/10, each narrowly rounded 120 apically and forming an inwardly projecting subapical tooth from the inner dorsal surface; median lobe (Figure 101) bearing paried, distally free serrated appendages ventrally, apex dorsally sagittate. Female: The female remains to be discovered. Distribution (Figure 145) CALIFORNIAN: CALIFORNIA: Siskiyou Co., Lake Mtn; lO-VI-l951; CASC (l 0' ). Trinity Co., Big Flat, Coffee Creek; 21-V1-1934; CASC (l o' ). Specimens examined: 2 db" . Diagnosis The serrate antennae, well developed tooth associated with the base of each tarsal claw, elytral apices which are flattened near each sutural margin but are neither impunctate nor shining, and parameres which are acutely rounded distally and which bear a well developed subapical tooth from the inner margin characterize the males of this extremely rare species. Remarks Several visits to the type locality and other known locations for this, the "rarest" North American species of Pedilus, proved fruitless as regards efforts to observe and collect additional material for study. Bionomics Nothing is known about the bionomics of dentatus. As a member of the punctulatus group, males would be expected to exhibit a positive orientation toward cantharidin but this has yet to be demonstrated. 121 5. Pedilus rustious Abrhllah (Figures 57, 72, 102, 146) Pedilus msticus Abdullah, l964b: 151, tb. 1, Figures 1-5. Type locality: "11 miles south of West Bakersfield, Kern County, California." Holotype,o‘ , BMNH. Pedilus inurbanus Abdullah, 1964b: 151, tb. 1, Figures 6-10. Type locality: "Coalinga, Jacalitos Canyon, Fresno County, California." Holotype, o' , BMN H. NEW SYNONYM. Description Male: Length 4-8 mm. Color of head scutellum, ventrally exposed portions of mesothorax and metathorax, legs, and abdominal sternites 3-8 black; prothorax orange to reddish orange. Head covered with moderately elongate erect setae, surface finely to moderately coarsely and sparsely to densely punctate, punctures most coarse and dense on, frons, between antennae and clypeus; antennae serrate. Pronotum ovate, covered with moderately elongate, retrorsely semierect setae, surface moderately coarsely and densely punctate. Legs with prothoracic and ' mesothoracic tarsi conspicuously tomentose beneath: tarsal claws each with a small, poorly developed basal tooth. Elytral surface moderately coarsely and densely punctate to rugose, lacking pruinose luster, entirely black or black with testaceous macula along sutural margin distally, macula attaining apex of elytra or not; elytral apices not modified. Abdomen with 7th sternite strongly deflexed, hind margin of 8th sternite broadly emarginate. Parameres (Figure 72) fused proximally, free and gradually divergent along distal 1/5, each obliquely 122 truncate apically, bearing a single, inwardly projecting subapical tooth from the inner surface: median lobe (Figure 102) bearing serrated appendages ventrally, apex slightly knobbed and strongly recurved, forming a ventral hook. Female: Differs from male by having tarsi of prothoracic and mesothoracic legs but slightly more tomentose beneath than those of metathoracic legs: elytra concolorom black, lacking testaceous sutural macula; hind margin of 7th abdominal sternite (Figure 57) conspicuously, acutely produced; 8th abdominal segment reduced, usually retracted within 7th, its posterior margin entire. Lateral extension of each ventral bacculus well developed. Distribution (Figure 146) CALIFORNIAN: CALIFORNIA: San Luis Obispo Co., 15 mi. SE. Simmler, S. end Soda Lake: 2-lV-1969; CASC (id). Fresno Co., Ciervo Hills, 13 air mi. sw Mendota; 16411-1975; CISC (3 99 ). Fresno Co., Ciervo Hills, 18 air mi. sw Mendota; 18411-1975; CISC (5 dd, 19 ). Fresno Co., Ciervo Hills, 18 air mi. SW Mendota; 19411-1978; DYCC (3 do' , 3 99 ). Fresno Co., Ciervo Hills, 18 air mi. sw Mendota: zs-m-lsvs; DYCC (3 0'0', 27 99 ). Fresno Co., Ciervo Hills, 18 air mi. sw Mendota; ls-Iv-lsso; DYCC, .1ch, MSUC (14 do‘ , 214 99 ). Fresno Co., Coalinga; 24-IV-1956; CDAE (2 99 ). Specimens examined: 26 dd, 250 99 . Diagnosis Males of rusticus exhibit the following diagnostic set of characters: frontoclypeal surface of head coarsely and densely punctate; elytral apices not modified; seventh abdominal sternite strongly deflexed ventrally; parameres obliquely truncate apically (Figure 72). Females are recognizable by the unique structure of the seventh abdominal sternite. The condition is approached in 123 cavatus, but the elytra of rusticus are black while those of cavatus are stramineous to testaceous with black apices, sutural margins and lateral margins. Remarks Having examined the holotypes of both rusticus and inurbamls, and having collected lengthy series from Fresno Co., CA, I am confident that only a single species is involved. The primary difference noted by Abdullah was that of punctation on the head. In many species of Pedilus, including msticus, punctation varies somewhat intraspecifically. Furthermore, within a single specimen punctation is commonly more coarse and dense toward the clypeus and less pronounced postocularly. The characters Abdullah used to distinguish between rusticus and inurbamls are well within the limits of intraspecific variability for most species of Pedilus. This study provides the first description of the female, easily identifiable by the characteristic form of the sevnth abdominal sternite. Bionomics Nearly all specimens of rusticus examined were collected at flowers of Phacelia tanacetifolia; two specimens were taken at Monolopia major, presumably on the flowers. The largest series, including 14 males and 214 females, was collected at the Ciervo Hills in western Fresno County, between 4:00-6:30 p.m. on 30 March 1980 (James Johnson, in litt.). Of the 228 specimens, 222 were taken at flowers of P. tanacetifolia, "6 from paler, flatter more open flowered Phacelia...." Although cantharidin bait was set out at each of the three recorded localities, no specimem oriented positively toward the compound. 124 6. Pedilus [W 81’. NOV. (Figures 73, 103-104, 147) Type Information Holotype: (0' ), 5 mi. N. Pearblossom, Lovejoy Buttes, L.A. Co., Calif.; IV- 12-64, R.L. Langston, Collector, [California Academy of Sciencesi. Paratopotypes: 2 o‘o' , same data as holotype, [l in CISC, l in DYCC]. Paratypes: 10' , Randsburg [Kern CoJ, Cal., T. Craig, 4-5-‘27, T. Craig Collection, [CASC]; 10‘ , Ft. Tejon [Kern Col, Cal., Iv-ls-zs,0' , H.C. FALL COLLECTION, [MCZC]. Description Male: Length 5—6 mm. Color of labial palpi and sclerotized portions of genitalia testaceous; prothorax orange; remaining visible regions of head, thorax including elytra, and abdomen rufopiceous to black. Head covered with moderately elongate semierect to erect setae; frontoclypeal surface finely and densely punctate, postocular aspect of genae and remainder of cranial surface finely, sparsely punctate; antennae serrate. Pronotum transversely ovate, its maximal width 3/2 X its mesal length, surface covered with moderately elongate, mostly retrorsely semierect setae, finely and very sparsely punctate. Legs with prothoracic and mesothoracic tarsi conspicuously dilated and tomentose beneath; tarsal claws each with a small, poorly developed basal tooth. Elytral surface covered with short retrorsely semierect setae, coarsely and densely punctate to rugose: elytral apices very weakly impressed. Abdomen with 7th sternite slightly deflexed, bearing a fringe 125 of prominent setae along the subtruncate hind margin; 8th sternite slightly emarginate along posterior margin. Parameres (Figure 73) fused proximally, free and slightly divergent along distal US, each forming an inwardly projecting subapical tooth from the inner dorsal surface; median lobe (Figures 103-104) bearing paired serrated appendages which are recessed within the excavated ventral surface, apex knobbed and recurved, forming a ventral hook. Female: The female remains to be discovered. Etymology In gratitude for his many kindnesses related to these studies, and for donations of the Pedilus specimens in his personal collection, this species is named for Mr. Kenneth M. Fender of McMinnville, Oregon. Distribution (Figure 147) CALIFORN'IAN: CA. See type information for specific localities. [CASC, CISC, DYCC, MCZC.] Specimens examined: 5 db' . Diagnosis Several western Species of Pedilus, including f enderi , have the seventh abdominal sternite of the male flanged or deflexed ventrally. The following set of characters should serve to separate f enderi from other members of this group: frontoclypeal surface of head finely and densely punctate; elytral apices weakly, - juxtasuturally impressed; parameres narrow, acutely rounded apically. 126 Remarks In late March, 1978, the type locality of this distinctive new species at Lovejoy Buttes was briefly visited. lnclimate weather and a paucity of flowering plants hampered collecting efforts and no additional specimens were found. It was, however, noted that much of this small and possibly unique habitat was being altered by construction of new homes. The extent and outcome of man's perturbation in the Lovejoy Buttes remain to be evaluated. Bionomics No information is available regarding the natural history of this species. However, Pedilus cavatus, one of its closest relatives, has been shown to be very strongly "attracted" to cantharidin and the same type of activity might be expected to hold for males of fenderi. 127 7. Pedilus flexiventris Fall (Figures 58, 74, 105, 148) Pedilus flexiventris Fall, 1915: 26-27, Figure 12; Leng, 1920: 161 (catalog); Abdullah, 1962: 219. Type locality: Pasadena, California. Holotype, o‘ , #24316, MCZC. Pedilus flexiventris (Fall); Abdullah, 1966b: 175-176, Figures 25-30 (distribution). Description Male: Length 5-8 mm. Color of head, scutellum, ventrally exposed portions of mesothorax and metathorax, and abdominal sternites 3-8 piceous to black; prothorax yellowish-orange to orange. Head covered with elongate semierect and erect setae, postocular aspect of genae finely and sparsely punctate. Antennae serrate, rufopiceous to black. Pronotum ovate, covered with moderately elongate setae, finely and sparsely punctate. Legs piceous to black; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws each with a small, poorly developed basal tooth. Elytra densely punctate to somewhat rugose, black with yellowish to stramineous apices, dark portions frequently with a pruinose luster; elytral apices deeply, obliquely impressed. Abdomen with 7th sternite strongly deflexed, bearing a fringe of prominent setae along hind margin; posterior margin of 8th sternite emarginate. Parameres (Figure 74) fused proximally, free and slightly divergent along distal US, each forming an inwardly projecting subapical tooth from the inner surface; median lobe (Figure 105) bearing paired, distally free serrated appendages which are recessed within the excavated 128 ventral surface, apex knobbed and recurved, forming a ventral hook. Female: As for male except tarsi of prothoracic and mesothoracic legs but slightly more tomentose beneath than those of metathoracic legs; elytral apices not impressed; 7th abdominal sternite (Figure 58) not at all deflexed; 8th abdominal segment usually telescoped within 7th, not visible. Lateral extension of each ventral bacculus well developed. Distribution (Figure 148) CALIFORNIAN: CA. [AMNH, CASC, CDAE, CISC, CNCI, CUIC, DEFW, DYCC, GHNC, INHS, LACM, MCZC, MSUC, OSUC, SEMC, UCRC, USNMJ Specimens examined: 51 0' 0' , 47 99 . Along the coast, most specimens have been collected between the last week of April and the last of May; this interval is delayed by about two weeks in the inland foothills and mountains. Diagnosis Males of flexiventris may be characterized as follows: elytra black with yellowish to stramineous apices; elytral apices abruptly, deeply obliquely impressed; seventh abdominal sternite strongly deflexed ventrad. Of the female Pedilus which have a small basal tooth associated with the tarsal claw, only flexiventris has the following set of characters: dorsal surface of head covered with moderately elongate yellowish setae; postocular surface of head finely, sparsely punctate;'head, scutellum and elytra black. Remarks Abdullah (1966b: 176) listed Utah, in addition to California, as a state from 129 which specimens of flexiventris were seen. This author has seen no material from Utah and doubts very much whether the species occurs outside the established range of southwestern California. Any substantial new records would most likely come from northern Mexico. Bionomics A single male from San Diego County bears the label "alfalfa." The close relationship of this species with cavatus indicates that positive cantharidin orientation may eventually be demonstrated for flexiventris males. 130 8. Pedlus cavatus Fall (Figmes 20, 54, 75, 106, 149) Pedilus cavatus Fall, 1915: 25-26, Figures 12, 12a; Leng, 1920: 161 (catalog); Abdullah, 1962: 219: Hatch, 1965: pl. 16, Figure 9, pl. 17, Figure 7. Type locality: Alameda Co., California. Holotype, o‘ , #24314, MCZC. Pedilus cavatus (Fall); Abdullah, 1966b: 174-175, Figures 19-24 (distribution). Description Male: Length 5-7.5 mm. Color of head, ventrally exposed portions of mesothorax and metathorax, and abdominal sternites 3-8 piceous to black; pronotum orange. Head densely covered with elongate semierect and erect setae, postocular aspect of genae finely and sparsely punctate; labrum sometimes testaceous to piceous; first 3 segments of maxillary palpi testaceous to black, terminal segment piceous to black. Antennae (Figure 20) serrate, rufopiceous to black. Pronotum ovate, densely covered with moderately elongate setae, finely and sparsely punctate; prosternum orange with black posterior margin; scutellum testaceous to black. Legs usually testaceous with femorotibial articulations and portions of tarsi piceous to black; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws each with a small, poorly developed basal tooth. Elytra finely and densely punctate, stramineous to testaceous with narrow black sutural and lateral margins; elytral apices deeply, obliquely impressed (Figure 54) with anterior aspect of impressed area black and apex yellowish to testaceous. Abdomen with 7 th sternite strongly deflexed, 131 bearing a fringe of prominent setae along hind margin; sternites 7 and 8 broadly emarginate along posterior margins. Parameres (Figure 75) fused proximally, free and slightly divergent along distal US, each forming an inwardly projecting subapical tooth from the inner dorsal surface; median lobe (Figure 106) bearing paried, distally free serrated appendages ventrally, apex knobbed and recurved, forming a ventral hook. Female: As for male except tarsi of prothoracic and mesothoracic legs but slightly more tomentose beneath than those of metathoracic legs; elytral apices not impressed; abdominal sternite 7 not at all deflexed, its hind margin rounded, 8th abdominal segment telescoped within 7th, usually not visible. Lateral extension of each ventral bacculus well developed. Distribution (Figure 149) PACIFIC NORTHWEST: OR, WA; CALIFORNIAN: CA. [AMNH, BMUW, CASC, CDAE, CISC, CNCI, DYCC, PMNH, JBJC, JSCC, LACM, MCZC, MSUC, OSUO, UCDC, USNM, WSUCJ Specimens examined: 526 0‘0” , 32 99 . Best collecting for cavatus has been between the last week of April and the end of May. Diagnosis The abrupt, deep, oblique impression of the elytral apices and strong ventral deflexion of the seventh abdominal sternite serve to separate males of cavatus from all but those of flexiventris The elytra of flexiventris are black with yellowish to stramineous apices while those of cavatus are stramineous to testaceous with black apices, sutural margins and lateral margins. Females of 132 cavatus may be characterized as follows: dorsal surface of head densely covered with elongate yellowish-gold setae; basal tooth of tarsal claw small; elytral color similar to that of male. Remarks This species was poorly represented in collections at the onset of this study. Fall (1915) based his description on 15 specimens, Abdullah (1966b) examined only 60 specimens and a mere 68 were received on loan. However, present collecting efforts, particularly with the utilization of cantharidin bait, has dramatically increased the number of individuals available for study. The specimens from Oregon and Washington represent new state records for cavatus. Previously, it was known only from the coastal and Sierra-Nevada Mountain regions of central and northern California. Bionomics Pedilus cavatus has been collected on the legume, Prosopis in Contra Costa County, on Azalea flowers in Napa County, from foliage of oak in Marin and Solano counties, and from foliage of Hippocastaneum and U mbellularia and an erect, blue-flowered C eanothus bush, all from Gates Canyon in Solano County. Hundreds of males and three females have been collected at cantharidin bait from early April to early June in Butte, Humboldt, Marin, San Mateo, Santa Cruz, Solano, Trinity, and Yolo counties. Stream margins and edges of wooded areas have been the most productive areas for placement of baits. 133 9. Pedilus jot-mun SP. NOV. (Figures 59, 76, 107-108, 150) Type Information Holotype: (0' ), CA: Kern County, Devils Den Road, 1.5 mi W Hwy 33, 26- 111-1978, COLLECTED BY: James Johnson, Janice Johnson, Elev 600 feet, taken at cantharidin bait [California Academy of Scienced. Allotype: ( 9 ), same data as holotype except, Ex: Flowers of Mon010pia lanceolata, [CASC]. Paratopotypes: 3 0' 0' , 20 99 as follows: ld' , 7 99 , same data as allotype; 7 99 , same data as allotype except, 20-111-1978, COLLECTED BY: James Johnson, Janice Johnson, Daniel Young, Joan Young; 4 99 , same data as allotype except, 29—30 Mar. 1978, [2 collected by] J.T. Johnson, [2 collected by] J.B. Johnson; 1 9 , same data as allotype except, 29-30 Mar. 1978, J.B. Johnson, taken at cantharidin bait; l 9 , same data as allotype except, 29—30 Mar. 1978, J.T. Johnson, taken from flowers of PHACELIA SP.; 2 o'o' , CA: Kern County, Devils Den Road, 1.5 mi W Hwy 33, 16-IV-l980, C.Y. Kitayama, collector, in flight [caJ 10:00 a.m. Paratypes: 11 do" , 12 92 as follows: 10 do' , 12 99 , CAL. San Luis Obispo Co. S. end Soda Lake, 15 mi SE. Simmler, 2.1V.l969, Hugh B. Leech coll.; lo' , CALIF: Kern Co., 2 mi W Devil's Den, lll-l9-75, J.A. Powell. Paratypes are deposited in the following collections: BMN H, CASC, CISC, DYCC, JBJC, MCZC, MSUC, and USNM. 134 Desaiption Male: Length 4.5—7.5 mm. Color of head, ventrally exposed portions of mesothorax and metathorax, legs, elytra and abdominal sternites 3-8 black; pronotum orange to reddish—orange; prosternum usually black, occasionally black with anterior portion orange to reddish-orange. Head covered with moderately elongate erect setae, surface moderately coarsely and densely punctate. Antennae serrate, articles 1-2 frequently more darkly pigmented and shining than remaining segments. Pronotum ovate, covered with erect and retrorsely semierect setae, surface moderately coarsely and densely punctate. Prothoracic and mesothoracic tarsi but slightly more expanded and tomentose beneath than those of metathoracic legs; tarsal claws each with a small, poorly developed basal tooth. Elytral surface moderately coarsely and densely punctate to rugose, lacldng pruinose luster; elytral apices not modified. Abdomen with hind margin of 7th sternite entire, nearly straight, distal margin of 8th sternite broadly, shallowly emarginate. Parameres (Figures 76, 108) fused proximally, free and subparallel along distal 3/13, acutely rounded apically, each bearing a single, well developed dorsoanteriorly directed subapical tooth from the inner ventral surface; median lobe (Figure 107) bearing paired, distally free serrated appendages which are partially recessed within the excavated ventral surface, lateral aspects subparallel along most of their length, rather abruptly converging distally, thus forming a blunt point. Female: As for male except color far more variable: head, scutellum, ventrally exposed portions of thorax, coxae, trochanters and femora orange to reddish-orange, reddish—orange suffused with black, or entirely black; elytral epipleura and lateral margins also commonly reddish—orange. Antennae weakly 135 serrate, posterior margin of 8th abdominal sternite nearly entire, very shallowly emarginate in a few specimens. Lateral extension of each ventral bacculus well developed. Etymology I am very pleased to name this species for my dear friends, "Ding" and Jan Johnson. They spent many hours in the field attempting to collect specimens of. johnsonorum as well as numerous other species of western Pedilus. The data base for this study has been significantly enhanced by the contributions of material they have made. Distribution (Figure 150) CALIFORNIAN: CA. See type information for specific localities. [BMNH, CASC, CISC, DYCC, JBJC, MCZC, MSUC, USNM.] Specimens examined: 1360' , 33 99 . Diagnoa'n The following set of characters should separate males of johnsonorum from all other North American Pedilus males: antennae serrate; elytral apices not modified; apex of median lobe produced into an acute spine-like process (Figure 107); parameres each bearing a well deve10ped dorsoanteriorly directed subapical tooth from the inner ventral surface (Figures 76, 108). The females are identifiable by the color of the head and scutellum (orange to reddish-orange or orange suffused with black) and by the small tooth associated with the base of each tarsal claw. 136 Remarks Although color variation was observed to be far greater in females, there does appear to be close correlation within the females examined relative to pigmentation of the head, thorax and legs (e.g. specimens with reddish-orange heads also tend to have similarly colored thoraces and legs). Bionomics Both the type locality near Devil's Den and the other known site for this species near the south end of Soda Lake are characterized by open grassy hills. Although most slopes were dominated by California Poppy during the author's visit to the type locality, most specimens of johnsonorum were observed feeding at flowers of the composite, Monolopia lanceolata. A single female was also taken at flowers of an unidentified species of Phacelia, and single specimens of each sex were collected from cantharidin bait on separate visits. The Devil's Den locality was revisited on 30 March and once again on 16 April, 1980 (James Johnson, in litt.). Very few poppies and no Monolopia were noted and not a single johnsonorum was observed. A sign warning of "PCP contamination" was pointing in the direction of the hill which produced all specimens collected in 1978; the extent and implications of the contamination on this habitat and population of johnsonorum are unknown. Phoretic soproglyphid deutonymph mites of the genus Calvalia were attached to several specimens of johnsonorum from the type locality (kindly identified by W. Calvin Welbourn; specimens deposited in acarology collection, Ohio State University, and the author's personal collection). According to Welbourn (in littJ, "adults of these mites are usually associated with wet vegetable matter." 137 Since it is unlikely that the mites attach to johnsonorum on flowers exposed to the sun and wind, the following scenerio is hypothesized: johnsonorum larvae are associated with moist decaying vegetative materials on or just below the soil surface (e.g. roots and prostrate stems of dead Monolopia, etc.). Mites become attached to adult jolmsonorum soon after the beetles emerge from the pupa, perhaps while they are still teneral and relatively vulnerable to "attack." 138 10. Pedilus inconspiculs (Horn) (Figures 21, 46, 60, 77, 109-110, 151) Corphyra inconspicua Horn, 1874: 42; Horn, 1883: 310, Figure 14. Type locality: "California." Lectotype and paralectotype designated by Abdullah, 1962: 219; lectotype, o‘ , #3038, MCZC; paralectotype, o' , #7985, MCZC. Pedilus (Corphyra) inconspicuus Horn; Pic, 1911: 13 (catalog). Pedilus inconspicuus Horn; Fall, 1915: 28-29, Figure 14 (distribution). Pedilus inconspicuus var. flavidus Fall, 1915: 28-29; Abdullah, 1962: 219. Type locality: El Dorado Co., California (from specimen label). Holotype, o' , #24315, MCZC. NEW SYNONYM. Pedilus inconspicuus (Horn); Leng, 1920: 162 (catalog); Abdullah, 1962: 219 (designation of lectotype); Abdullah, l964d: 13-15, Figures 7-12, map 1 (distribution). Pedilus inconspicuus flavidus Fall; Leng, 1920: 162 (catalog). Pedilus flavidus Fall; Tanner, 1927: 25, Figures 72-73 (female external genitalia); Abdullah, 1964d: 15-18, Figures 13-18, map 2 (distribution, bionomics). Pedilus pratti Hatch, 1965: 119. Type locality: "Coupeville, Wash., Sunnyside." Holotype, 0' , OSUO [holotype not examined. NEW SYNONYM. Type Information Although Horn listed only "California" as the type locality for inconspicua, Fall (1915: 28-29) stated, "...there is little doubt that they [= the syntype series] were taken somewhere in Middle California." The specimen designated to serve as the lectotype is from the Horn collection; the paralectotype, referred to by 139 Abdullah (1962: 219) as a "paratype," is in LeConte's collection; both agree with Horn's original description. Description Male: Length 4-8.5 mm. Color of head, scutellum, ventrally exposed portions of mesothorax and metathorax, legs and abdominal sternites 3-8 piceous to black; pronotum yellowish-orange to reddish—orange, rarely with disk piceous; prosternum yellowish-orange to reddish-orange, commonly with hind margin piceous to black; elytra entirely piceous to black or testaceous with variable amounts of light brown to black coloration along sutural and lateral margins, and apices. Head covered with moderately elongate, retrorsely semierect setae, surface moderately coarsely to coarsely and moderately densely punctate, mouthparts, labrum and anterior portion of clypeus testaceous to black. Antennae (Figure 21) serrate, apices of articles 1-2 testaceous to rufotestaceous, basal portions of segments 1-2 and remaining articles piceous to black. Pronotum ovate, covered with moderately elongate, retrorsely semierect setae, surface moderately finely but moderately densely punctate. Prothoracic and mesothoracic tarsi but slightly more tomentose beneath than those of metathoracic legs; tarsal claws (Figure 46) each with a small, poorly developed basal tooth. Elytral surface moderately coarsely and densely punctate to rugose, lacking pruinose luster; elytral apices not modified. Abdomen with posterior margins of sternites 7-8 broadly emarginate. Parameres (Figures 77, 110) fused proximally, free and subparallel along distal 3/10, each rounded apicany and bearing an inwardly projecting, serrulate, molar-like process from the inner 140 surface; median lobe (Figure 109) bearing paired, distally free serrated appendages which are recessed within the excavated ventral surface, lateral aspects of median lobe subparallel along most of their length, abruptly convergent distally, thus forming a stout apical tooth. Female: As for male except posterior margins of abdominal sternites 7 (Figure 60) and 8 entire. Lateral extension of each ventral bacculus well developed. Distribution (Figure 151) PACIFIC NORTHWEST: BC, OR, WA; CALIFORNIAN: CA. [AMNH, BMNH, BMUW, CASC, CCNY, CDAE, CISC, CUIC, DEFW, DSZI, DYCC, FMNH, GHNC, JBJC, JSCC, LACM, MSUC, NAUF, NMPC, OADC, OSUC, OSUO, UCDC, UCRC, UMRM, USNM, WSUCJ Specimens examined: 878 o'o' , 158 99 . This species has been collected in large numbers between late March and the middle of May along the coast and a 10-14 days later in the inland counties. Diagnosis Males of inconspicluls may be characterized as follows: antennae serrate; elytral apices not modified; apex of median lobe produced into an acute mesal tooth-like process (Figure 109); parameres each bearing an inwardly projecting, serrulate, molar-like process (Figures 77, 110). Of the female Pedilus which have a small basal tooth associated with the tarsal claw, only inconspicuus has the following combination of characters: dorsal surface of head with punctation somewhat variable, postocular aspect very coarsely and densely punctate; head sparsely covered with moderately elongate setae; head and scutellum piceous to black. 141 Remarks The primary source of intraspecific variation resides in elytral pigmentation. This prompted Fall (1915: 28) to describe the "variety," flavidus: For the sake of convenience, this name is proposed for a form of the above species [i.e., inconspicuusl having the elytra rufotestaceous or yellow with the sutural edge, side margin, at least toward the apex, and the tip blackish. Of the two forms, Fall wrote: The black form occurring in the San Francisco Bay region may therefore, I think, be safely regarded as typical. Along with them, and even more abundant, judging from the material at hand is found the pale form—var. flavidls. Farther east, in the foothills of the Sierras, from Tehama to Tulare Co., the flavidus form alone prevails, at least there are no black examples from this region in the material examined. Farther to the North—in Siskiyou Co. the black form and a dusky pallescent variety of it occurs, while throughout Southern California the black form alone is found. In 1964, Abdullah (l964d: 15) elevated flavidus to species level, largely on the basis of color differences. Having examined over a thousand specimens of this taxon, no evidence in terms of color or any other character discontinuity was found to warrant the recognition of two species. If anything, the character analysis suggests a continuous, North-South cline with respect to elytral pigmentation (Table 16). Seven paratypes of Pedilus pratti Hatch (six from the JSCC and one from the OSUO) were also examined; it is, without doubt, conspecific with inconspicwls. In addition to British Columbia, Washington, Oregon and California, Abdullah also reported examining specimens from Alberta, Montana, Nevada and Arizona. Since not a single specimen was seen to confirm any of these records, they have not been included with the distributional data. Table 16. Pedilus inconspicuus (Horn): 142 Analysis of Elytral Color Morphs in Western North America (B - Black Elytra; Y - Pale Form). #BzY 2 "Black" Oregon 11:117 91 Northern CA 11:100 102 Central CA, 108:427 20% Southern CA 56:53 51% 143 Bionomics Abdullah (l964d) communicated the following plant associations: on Pinus ponderosa, on Quercus agrifolia, on Q. lobata, and on Papulus trichocarpa. The large number of specimens of this common species have provided numerous additional records. In Oregon, inconspicwls has been collected by "sweeping" and "sweeping grass," at "Mech. trap," and at "choke cherry." Two specimens were taken at a malaise trap baited with dry ice in Marin County, others were labelled "sweeping" or "swept from grasses" in Alameda, Contra Costa and Tulare counties, all in California. Other California collectiom note the following: "rotary trap" and Prunus subcordata, Tulare County; Ceanothus, Tuolumne County; Pinus ponderosa and Ceanothus integerrimus, El Dorado County; Cerocarpus betuloides, N apa County; Rhus diversiloba and Rammculus, Santa Clara County; Heracleum, Contra Costa County; Lomatium, Monterey County; Monolopia major, Fresno County; "Morning Glory," Santa Barbara County; Rhus ovata, "beating Ceanothus tomentosus," and "collected from Pea," Riverside County. Large numbers of male inconspicuus were collected at cantharidin bait near McMinnville, Oregon, and at numerous sites in California. One sample from near Somerset, El Dorado County, contained several female inconspicuus in addition to males. An analysis of material collected at cantharidin bait led Robert Wharton to suggest (in litt), that this species "apparently prefers the open fields" at the Somerset location in deference to nearby grassy, shaded riparian habitats. On one occasion, specimens at cantharidin bait were observed being preyed upon by an unidentified lizard (James Johnson, in litt). 144 Larvae of this species were reared from decaying acorns of Quercus kelloggi Newberry at the Somerset location (Wharton, 1979). Acorns occupied by inconspicuus larvae were found beneath a large Q. kelloggi in an open field at depths of 5-10 cm. below the soil surface. 145 ll. Pedilus vittatus (Rom) (Figures 51, 64, 78, 111, 152) Corphyra vittata Horn, 1871: 279; Horn, 1874: 41; Horn, 1883: 310, Figure 15. Type locality: "Amador Valley" [California]. Holotype,0' , #3040, MCZC. Pedilus (Corphyra) vittatus Horn; Pic, 1911: 14 (catalog). Pedilus vittatus Horn; Fall, 1915: 30-31, Figure 11 (distribution). Pedilus lineatus Fall, 1915: 31, Figure 11; Leng, 1920: 162 (catalog); Abdullah, l964d: 23-24, Figures 37-38, map 4 (distribution). Type locality: Pasadena, California. Holotype, 0' , #24318, MCZM. NEW SYNONYI. Pedilus vittatus (Horn); Leng, 1920: 162 (catalog); Abdullah, 1962: 219; Abdullah, 1964d: 23, Figures 31-36, map 5 (distribution). Description Male: Length 6-9 mm. Head and ventrally exposed portions of mesothorax and metathorax black; mouthparts, antennae and scutellum testaceous to piceous; pronotum reddish-orange to rufopiceous, occasionally variously suffused, usually with disk darkest; prosternum yellowish-orange to testaceom anteriorly, rufopiceous to black posteriorly. Head covered with moderately elongate, retrorsely semierect setae, surface coarsely and densely punctate; antennae serrate. Pronotum ovate, covered with moderately elongate, retrorsely semierect setae, surface coarsely and densely punctate. Color of legs highly variable, testaceous suffused with darker pigmentation at femorotibial articulations to entirely black; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws each 146 bearing a small basal tooth. Elytral surface finely and densely punctate to rugose, commonly with pruinose luster; each elytron frequently rufopiceous to black with a testaceous vitta, broadest near humeral angle, terminating preapically; many specimens with elytra concolorous testaceous; elytral apices not modified. Venter of abdomen testaceous to black, commonly rufopiceous; hind margin of 7th sternite subtruncate, that of 8th sternite broadly, shallowly emarginate. Parameres (Figure 78) fused proximally, free and subparallel along distal US, each obtusely rounded apically and bearing a blunt, inwardly directed subapical tooth from inner surface; median lobe (Figure 111) bearing paired, distally free, serrated appendages ventrally and phallic papillae ventromesally, apex tapering to an acutely rounded point. Female: As for male except tarsi of prothoracic and mesothoracic legs but slightly more tomentose beneath than those of metathoracic legs; posterior margin of 8th abdominal sternite entire, the 8th segment usually retracted within 7th. Lateral extension of each ventral bacculus (Figure 64) strongly reduced, visible only as a thickening of the bacculus. Distribution (Figure 152) MOUNTAIN: UT; PACIFIC NORTHWEST: ID, OR; CALIFORNIAN: CA. [AMNH, BMNH, BMUW, CASC, CDAE, CISC, CNCI, DCCC, DEFW, DYCC, EMUS, FMNH, MCZC, MSUC, UCDCJ Specimens examined: 16 do' , 71 99 . Along the coast, most collections were made between late April and the second week of May. The best time to find vittatus adults in the inland areas is between the third week of May and the third week of June. 147 Diagnosis Like bardi, oregonus and punctulatus, males and females of vittatus have serrate antennae, a small basal tooth associated with the tarsal claw and coarse, dense punctation associated with the dorsal surface of the head and pronotum. Males of vittatus differ from these species by the structure of the elytral apices: caudate and upcurved in bardi, widely flattened along the distolateral margins in oregonus, impunctate and somewhat swollen in ptmctulatus, and not modified in vittatus. In the females of bardi, oregonus and MCMGUJS, the lateral extension of the ventral bacculus is rod—like; it is reduced and visible only as a lightly sclerotized baccular plate in female vittatus. Remarks No evidence was found to support recognition of the species ”"90th proposed by Fall (1915) and followed by Abdullah (1964d). The color and punctation characters these authors alluded to for differentiating between vittatus and lineatus are the same features which have been shown time and again to be far too variable for recognition of most species of Pedilus. The only additional character mentioned was the mesal impressed line associated with the pronotum of lineatus. However, even Fall and Abdullah noted that in the material they attributed to lineatus the line was of variable length, while some vittatus also possessed the sulcus. The Utah specimens represent a new state record; Abdullah (l964d) also recorded vittatus from British Columbia. 148 Bionomics Horning and Barr (1970) listed this species from Balsamorhiza sagittata and H eracleum lanatum at Craters of the Moon National Monument in ldaho; a single specimen was examined which was taken on Senecio serra at the same locality. At Marin County, California, one specimen was recovered from a Berlese sample of a "Neotoma debris pile." Associated with three Alameda County, California, specimens was the bionomics label, "among roots." According to the collector, these specimens were reared from larvae which were found tunnelling in the root system of the nettle, Urtica dioica (Wayne Gagne, in litt.). 149 12. PedilusaregamsFall (Figures 79, 112, 153) Pedilus oregonus Fan, 1915: 24-25, Figure 11; Leng, 1920: 161 (catalog): Abdullah, 1962: 219; Abdullah, l964d: 24-25, Figures 39—44, map 6 (distribution). Type locality: "Josephine Co., Oregon." Holotype, o‘ , #24320, MCZC. Description Male: Length 5.5-9 mm. Head black with palpi, labrum and anterior portion of clypeus testaceous to piceous, covered with moderately elongate erect and retrorsely semierect setae, surface coarsely and densely punctate. Antennae serrate, articles 1-2 often testaceous apically, remainder rufopiceous to black. Pronotum ovate, concolorous yellowish-orange to reddish-orange, rufopiceous or black, covered with moderately elongate decumbent setae, surface moderately coarsely and densely punctate; prosternum black, or orange anteriorly and black posteriorly; scutellum and ventrally exposed portions of mesothorax and metathorax piceous to black. Legs piceous to black, sometimes with variable testaceous regions; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws each bearing a poorly developed basal tooth. Elytra finely and densely punctate, commonly black with testaceous apices, occasionally concolorous testaceous or black, rarely black with indistinct testaceous vitta which is broadest near humeral angles, ending subapically; elytra widely flattened distolaterally. Venter of abdomen piceous to black; posterior margin of 8th 150 sternite emarginate. Parameres (Figure 79) fused proximally, free and slightly divergent along distal 3/22, each rounded apically and strongly toothed on inner margin; median lobe (Figure 112) bearing paired, distally free appendages ventrally, apex tapering to a point. Female: As for male except tarsi of prothoracic and mesothoracic legs but slightly more tomentose beneath than those of metathoracic legs; elytra usually concolorous black or testaceous; elytral apices not modified; hind margin of 8th abdominal sternite entire. Lateral extension of each ventral bacculus well developed. Distribution (Figure 153) PACIFIC NORTHWEST: BC, OR, WA; CALIFORNIAN: CA. [AMNH, BMUW, CASC, CISC, CNCI, DEFW, DYCC, PMNH, GHNC, JBJC, JSCC, MCZC, MSUC, NMDC, ODAC, osuo, SMCC, UCDC.) Specimens examined: 103 0'0‘ , 12 99 . The four week interval between the middle of May and June appears to be the best time to collect adults of this Species. Diagnosis The serrate antennae, small tooth associated with the base of each tarsal claw and coarse, dense punctation associated with the dorsal surface of the head and pronotum serve to separate males and females of oregonus from all but those of bardi, punctulatus and vittatus. Males of oregonus have the distolateral margins of the elytra widely flattened; the elytral apices of bardi are caudate and upcurved, those of punctulatus are impunctate and somewhat swollen, and those of vittatus are not modified at all. The rod-like lateral extension of each 151 ventral bacculus is common to females of bardi, oregonus and pimctulatus. The geographical range of bardi (southwestern California, Arizona) is significantly different from that of oregonus (northern California to British Columbia), but that of punctulatus overlaps with both. Thus, females of oregonus and punctulatus remain indistinguishable at this time. Remarlm . lntraspecific variation iS associated primarily with the coloration of the pronotum and elytra. The most common color pattern is orange pronotum and black elytra with yellowish-testaceous apices; all of the "yellowish-elytra" specimens examined also have an orange or yellowish-orange pronotum. The southern limit of this species appears to be in northern California. Extensive collections from numerous localities in California support this, and I must, therefore, take exception to the geographical distribution indicated by Abdullah (l964d: Map 6) in which the Species is illustrated as occurring as far south as Los Angeles County, California. Several specimens of punctulatus were examined which had been labeled as oregonus by Abdullah; this might account for his highly inaccurate distribution map. Bionomics A dozen specimens collected at Forest Grove, Oregon, in May and June of 1938 bear the label "mech. trap." Plant associations include "sweeping grass" and "on vetch" at Corvallis and Salem, Oregon, respectively. Males of oregonus have been observed and collected at cantharidin bait in Corvallis by Paul Johnson and in McMinnville, Oregon, by Kenneth Fender. 152 13. Pedilus bird! (Horn) (Figures 50, 80, 113, 154) Corphyra bardii Horn, 1874: 42. Type locality: San Buenaventura, California. Lectotype, designated by Abdullah, 1962: 219, o' , #3033, MCZC. Corphyra distinguenda Horn, 1874: 42; Horn, 1883: 307, Figure 15 (distribution). Type locality: San Buenaventura, California. Lectotype, designated by Abdullah, 1962: 219, 0', #3035, MCZC. NEW SYNONYI. Corphyra bardi Horn; Horn, 1883: 307, pl. 9, Figure 15. Pedilus (Corphyra) bairdi Horn; Pic, 1911: 12 (catalog; incorrect subsequent spelling)- Pedilus (Corphyra) distinguendus Horn; Pic, 1911: 12 (catalog). Pedilus bardii Horn; Fall, 1915: 22, Figures 6c, 11 (distribution). Pedilus bardii var. distinguendus (Horn); Fall, 1915: 22-23 (distribution). Pedilus bardi (Horn); Leng, 1920: 161 (catalog); Abdullah, 1962: 219 (designation of lectotype). Pedilus bardi distinguendus (Horn); Leng, 1920: 161 (catalog). Pedilus distinguendus (Horn); Abdullah, 1962: 219 (designation of lectotype); Abdullah, 1966b: 171-172, 174, Figures 13-18 (distribution). Pedilus bardii (Horn); Abdullah, 1966b: 170-171, Figures 7-12 (distribution). Description Male: Length 4.5-8 mm. Color of head and ventrally exposed portions of mesothorax and metathorax piceous to black, pronotum yellowish-orange to reddish—orange, other regions of body highly variable in color. 153 Head covered with moderately elongate erect setae, postocular aspect of genae coarsely and densely punctate. Antennae serrate, usually piceous, less frequently testaceous. Pronotum ovate, covered with moderately elongate erect to retrorsely semierect setae, coarsely and densely punctate; prosternum usually piceous to black posteriorly reddish—orange anteriorly, less frequently concolorous reddish-orange; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws each bearing a small basal tooth. Elytra usually black with yellowish to testaceous apices, occasionally concolorous testaceous or black with a longitudinal testaceom vitta extending from the humeral angle to the apex; elytral surface densely punctate to rugose, slightly pruinose or, more frequently, completely lacking pruinose luster; elytral apices impunctate and usually somewhat swollen; elytra caudate and curved upwardly apically. Abdominal sternites 3-8 piceous to black, occasionally suffused with testaceous coloration, sternite 8 emarginate; sclerotized portions of abdominal segments 9- 10 and genitalia testaceous to piceous. Parameres (Figure 80) fused proximally, free and slightly divergent along distal 3/ 20, each rounded apically and strongly toothed subapically on inner margin; median lobe (Figure 113) bearing paired, distally free and movable serrated appendages ventrally, apex tapering to a point. Female: As for male except tarsi of prothoracic and mesothorcic legs but slightly more tomentose beneath than those of metathorcic legs; elytra usually concolorous black, occasionally testaceous with piceous to black sutural and apical margins; elytral apices (Figure 50) not modified, neither swollen nor caudate and curved upwardly. ’ Abdominal sternite 8 with hind margin entire; lateral extension of each ventral bacculus well developed. 154 Distribution (Figure 154) SOUTHWEST: AZ; CALIFORNIAN: CA. [AMCC, AMNH, CASC, CDAE, CISC, CNCI, DYCC,1NHS, JBJC, LACM, MCZC, MSUC, chc, SEMC, UCDC, UCRC, UGCA, UMMZJ Specimens examined: 130 do' , 43 99 . The majority of adult bardi were collected between the second week of April and the middle of May, Slightly later in the mountains above 5,000 feet in elevation. Diagnosis Strongly caudate, upcurved elytral apices are unique to males of bardi. Females of bardi differ from all. but those of oregonus and punctulatus in the following character set: dorsal surface of head and pronotum coarsely and densely punctate; pronotum ovate, widest at the middle; basal tooth of each tarsal claw small; lateral extension of each ventral bacculus rod-like. The geographical range of bardi (southwestern California, Arizona) differs significantly from that of oregonus (northern California to British Columbia), but a reliable means for separating females of bardi from those of punctulatus has not yet been discovered. Remarks In addition to the obvious sexual difference associated with the elytral apices, intraspecific character variation is also manifest in elytral coloration. Black elytra ( 99 ) or black with yellowish apices ( 0'0' ) are by far the most . commonly encountered types. Of the 180 examined, 14 exhibited testaceous elytra while the elytra of an additional five specimens are black with a testaceous vitta. Only two of the 48 99 deviate from the typical condition; 155 both of them have testaceous elytra. Horn described both bardi and distinguendus from Specimens collected by Crotch at the same time and place. The two are clearly color variations of the same species, as Fall attempted to demonstrate when he considered distinguendls as a "variety" of bardi. I find no basis whatsoever for supporting the two species hypothesis which Abdullah resurrected. Bionomics Label data provide a number of plant associations for this species. In Ventura County, bardi was collected from the crucifer, Descurainia sophia and from "mixed grass lawn." Specimens were also taken on Salix and Pinus attenuata in Monterey County, from Senecio flowers in Orange County, by "sweeping grass under Quercus sp." on Santa Cruz Island, and from "sweet pea" in Santa Barbara County. The author and his wife collected bardi from flowers of Achillea millefolium var. lanulosa and Prunus Sp. which were growing near a stream in the Tehachapi Mountains of Kern County. Drier slopes a Short distance from the stream were beset with several species of flowering composites, but no bardi were observed. A Single male was collected "at light" in San Luis Obispo County. The first cantharidin record was provided by a male which was found in Fresno County "on Short, dry grass attacking and feeding on Colospasta elegans," a common meloid currently assigned to the genus Eupompha. Over 50 males were taken at cantharidin baits in Kern, Orange, and Santa Barbara counties during the course of this study; the Kern County Specimens were collected from bait at the edge of a wooded area along a stream. 156 14. PediluspuncmlatusLeConte (Figures 52, 81, 114-115, 155) Pedilus pimctulatus LeConte, 1851: 151; LeConte, 1855: 272; Fall, 1915: 23-24, Figures 6c, 11 (distribution); Abdullah, 1962: 219; Abdullah, 19661); 164, 166-170, Figures 1-6. Type locality: San Francisco, California (See Type Information, below, for discussion regarding disposition of "type.") Corphyra ptmctulata (LeConte); LeConte, 1867a: 65 (species list). Corphyra punctulata LeConte; Horn, 1871: 281-282; Horn, 1874: 41; Horn, 1883: 303, Figure 15. Corphyra funebris Horn, 1871: 280; Horn, 1874: 41; Horn, 1883: 307, Figure 15. Type locality: "California." Lectotype and paralectotype designated by Abdullah, 1962: 219: lectotype, 9 , #3037, MCZC; paralectotype, 9 , #8177, MCZC. Pedilus (Corphyra) funebris Horn; Pic, 1911: 12 (catalog). Pedilus (Corphyra) punctulatus LeConte; Pic, 1911: 13 (catalog). Pedilus punctulatus var. funebris Horn; Fall, 1915: 24. Pedilus punctulatus (LeConte); Leng, 1920: 161 (catalog). Pedilus punctulatus funebris (Horn); Leng, 1920: 161 (catalog). Pedilus fwtebris (Horn); Abdullah, 1962: 219 (designation of lectotype). Type Information It is clear from LeConte's description of punctulatus that both sexes were known to him, however, no single "type" was alluded to. Thus, Abdullah's (1962: 219) reference to a specimen in LeConte's collection as the holotype ( 9 , #4875) 157 was in error. It is possible that the specimen, the first in a series of 7-8 from California (gold disk is missing from 6th specimen), is from the syntype series. However, in view of curatorial inconsistencies over the years, any reference to specimens in either the LeConte or Horn collections must be made very carefully and thoughtfully (Newton, personal communication). The female specimen in question bears a "Pedilus punctulatus LeC. S.F." label in addition to the gold disk and "type" number label. On the same grounds as stated for ptmctulatus, no evidence was found to support Abdullah's lectotype and paralectotype designations for funebris. Description Male: Length 5-9 mm. Head and ventrally exposed portions of mesothorx and metathorax piceous to black; remainder of body highly variable in color. Head covered with short, retrorsely semierect setae, surface coarsely and densely punctate; mouthparts, labrum and anterior portion of clypeus testaceous to black. Antennae serrate, articles 1-2 testaceous to black, remaining segments rufopiceous to black. Prothorax with pronotum ovate, concolorous yellowish- orange to reddish-orange, variously suffused with piceous to black, or entirely black, covered with retrorsely semierect setae, surface moderately coarsely and densely punctate; color variation of prosternum similar to pronotum; scutellum piceous to black. Legs entirely testaceous, variously suffused with piceous to black pigm entation; or concolorous black; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws each bearing a small, (poorly developed basal tooth. Elytra moderately coarsely and densely punctate, entirely testaceous, variously suffused with piceous to black pigmentation, 158 testaceom with piceous to black sutural vitta and creamy yellowish to testaceous apices, or black with testaceous apices; elytral surface commonly with pruinose luster; elytral apices (Figure 52) impunctate and somewhat swollen, each tapering to a blunt point. Abdomen entirely piceous to black or piceous to black with variable yellowish-brown maculations, especially along lateral margins; 7 th sternite with hind margin entire, that of 8th sternite broadly emarginate. Parameres (Figure 81) fused proximally, free and subparallel along distal 3/ 20, each rounded apically and strongly toothed subapically on inner margin; median lobe (Figure 114) bearing paired, distally free, serrated appendages ventrally, phallic papillae and pectinations (Figure 115) ventromesally, apex tapering to an acutely rounded point. Female: Differs from male by having tarsi of prothoracic and mesothoracic legs but slightly more tomentose beneath than those of metathoracic legs; elytral apices neither swollen nor impunctate; abdominal sternite 8 with hind margin entire. Lateral extension of each ventral bacculus well developed. Distribution (Figure 155) PACIFIC NORTHWEST: OR, WA; CALIFORNIAN: CA. [AMNH, BMNH, BMUM, CASC, CDAE, CISC, CNCI, CUIC, DBUM, DEFW, DYCC, EMUS, FMNH, GHNC, INHS, JBJC, JSCC, LACM, MCZC, MSUC, NDSU, NMDC, NMPC, OSUC, PADA, PURC, SBMN, UASM, UCDC, UCRC, UMMZ, UMRM, USNM, VPIC, WSUCJ Specimens examined: 836 do" ,138 99 . Adults of punctulatus are most active during peak flowering times of such umbelliferous genera as Conium and H eracleum. 159 Diagnosis The following character set separates males and females of punctulatus from all but those of bardi, oregonus and vittatus: serrate antennae; coarse, dense punctation associated with the dorsal surface of the head and pronotum; a small tooth associated with the base of each tarsal claw. Males of punctulatus have the elytral apices impunctate and swollen; they are strongly caudate and upcurved in bardi males, widely flattened along the distolateral margins in males of oregonus, and not modified at all in vittatus males. The females of punctulatus have a rod-like lateral extension of each ventral bacculus, as do those of bardi and oregonus. The geographical range of bardi (southwestern California, Arizona) differs significantly from that of oregonus (northern California to British Columbia), but that of punctulatus overlaps with both. Thus, females of punctulatus may be tentatively identified over part of their range, but are indistinguishable at present from those of bardi and oregonus in areas where they overlap. Remarks This Species is highly variable with respect to coloration, particularly that of the pronotum, legs and elytra. While no clinal or other obvious patterns were noted, several series from Washington and Oregon did stand out. In these, a large percentage of specimens possess testaceous elytra with a relatively well developed black sutural vitta, much like that of typical vittatus Specimens. Abdullah (1966b) also listed punctulatus from British Columbia, Arizona, Colorado and Idaho. The occurrence of this Species in southern British Columbia and western Idaho is highly likely, but the validity of the Arizona and Colorado records is seriously doubted. 160 Bionomics Horn (1871) reported that punctulatus was found on "various composite plants"; it was recorded on pine, poppy, willow, and Ceanothus by Abdullah (1966b). The following plant associations were indicated by bionomies labels attached to California specimens examined during present studies: foliage of Baccharis pilularis at Albany, Alameda County; from flowers of Conium maculatum and Heracleum lanatum in Marin County; "ex: Wild Oaks-weeds" and on Solanum in Monterey County; on Ceanothus at LaHonda, San Mateo County; "ex. Oak" and "on poppy" in Santa Clara County; on Artemesia in Solano County; on a leaf of Salix in Sonoma County; "on grass" and Conium maculatum in Trinity County; from Salix at Davis, Yolo County. Numerous males of punctulatus have been observed and collected at cantharidin bait, especially from grassy areas near water. According to label data, a Single Pedilus larva collected by Hugh Leech in Marin County, California, "Emerged from dead stems of Baccharis pilularis var. consanguinea (adults reared from dead Lupinus, same place)." Two punctulatus males were examined from the collection of the California Academy of Sciences which should represent those referred to above. They emerged from dead stems of Lupinus arboreus (under the name L. propinqwls) which were collected at Marin County on 12 April 1957. The number of other Species of Pedilus which are known to occur in the same area raises doubts as to the implied association. Suffice to say that the larva may be that of punctulatus. THE LEWISI GROUP Diagnosis The excurved parameral apex and the presence of a spine or tooth-like process associated with the ventral or ventrointerior aspect of each paramere (Figures 82-85) serve to characterize this group. Remark Three of the four Species comprising this group are western while the fourth, elegans, is a common eastern Species. 161 162 15. Pedilus picipeu‘tis Fall (Figures 61, 82, 116, 156) Pedilus picipennis Fall, 1915: 29-30, Figure 16; Leng, 1920: 162 (catalog): Abdullah, 1962: 219; Abdullah, l964f: 97-93, Figures 39-44, map 4 (distribution). Type locality: Lake Tahoe, California ("Tallac" according to specimen label). Holotype, 0' , #24322, MCZC. Description Male: Length 5-7 mm. Color of head, ventrally exposed portions of mesothorax and metathorax and abdominal sternites 3-8 piceous to black, mouthparts, antennae, scutellum and elytra testaceous to black; pronotum, usually rufopiceous to black, rarely orange; prosternum rufotestaceous, suffused variously with piceous to black pigmentation, or entirely black; legs testaceous, black or light brown variously suffused with black. Head covered with short, semierect setae, surface moderately coarsely and sparsely punctate; antennae serrate. Pronotum ovate, covered with retrorsely semierect setae, surface moderately coarsely and sparsely punctate. Legs with prothoracic and mesothoracic tarsi scarcely more dilated and tomentose beneath than those of metathoracic legs; tarsal claws each with a small, poorly deve10ped basal tooth. Elytral surface moderately coarsely and densely punctate to rugose, lacking pruinose luster; elytral apices not modified. Hind margins of abdominal . sternites 7-8 broadly emarginate; 8th segment often retracted within the 7th. Parameres (Figure 82) fused proximally, free and Slightly divergent along distal l/ 2, each rounded apically, excurved subapically forming a well developed, 163 outwardly projecting tooth, each bearing a broad, inwardly projecting subapical tooth from the inner margin; median lobe (Figure 116) with sides subparallel, slightly tapering apically, apex rounded. Female: Differs from male by having the hind margins of sternites 7 (Figure 61) and 8 broadly rounded, entire, 8th segment usually retracted within the 7 th. Lateral extension of each ventral bacculus well developed. Distribution (Figure 156) MOUNTAIN: UT; PACIFIC NORTHWEST: ID, OR, WA; CALIFORNIAN: CA. [AMNH, BMUW, CASC, CDAE, CISC, DYCC, FMNH, JBJC, JSCC, MSUC, NMDC, OSUC, OSUO, UCDCJ Specimens examined: 63 0'0' , 17 99 . The best time to collect picipennis adults is between the third week of June and the middle of July, 1-2 week later in the mountains above 5,000 feet in elevation. Diagnosis The small basal tooth associated with each tarsal claw, elytral apices which are neither impressed nor impunctate and swollen, and parameres which are excurved subapically serve to characterize males of picipennis The excurvate type of paramere is also found in males of elegans, granti and lewisi, but all three have the basal tooth of each tarsal claw well developed and elytral apices which are variously impressed, swollen surrounding the impressed area, and impunctate. Females of picipennis have the following diagnostic combination: head covered with short setae, postocular surface finely, sparsely punctate; head, scutellum and elytra testaceous to black; pronotum ovate; basal tooth of each tarsal claw reduced, small; seventh abdominal sternite broadly rounded posteriorly. 164 Remark The majority of specimens examined possess rufopiceous to black pronota and testaceous to brown elytra; in several, the elytra are black, while three other Specimens have orange pronota. Abdullah (1964f) listed the following additional localities which were not confirmed by specimens in this study: Alberta, Colorado and Montana. Bionomics Label data indicate that most PiCiPOMis have been collected at elevations of 5,000-9,000 feet. In El Dorado County, California, a single male was taken from flowers of Veratrum californicum growing in a montane meadow at 6,700 feet. Males from El Dorado, Mariposa and Tuolumne counties, California, were collected at cantharidin bait placed adjacent to streams running through grassy meadows sparsely forested with Populus tremuloides, P. trichocarpa, and Salir app- 165 16. Pedlus elegans (Bent!) (Figures 42, 64, 83, 117-118, 157) Pyrochroa ? elegans Hentz, 1830: 257. Type locality: "Massachusetts." Holotype, 9' (inferred from description), [not examined, repository unknown]. Pedilus fulvipes Newman, 1838: 375; LeConte, 1855: 274. Type locality: "N. A." Holotype, 9 , BMNH. Pedilus lugubris Newman, 1838: 375; LeConte, 1847: 84. Type locality: "N. A." Holotype, 0' , BMN H. Pedilus haemorrhoidalis Ziegler, 1844: 46—47. Type locality: "Pennsylvania." Holotype, 0' , MCZC. Pedilus ruficollis Ziegler, 1844: 47; LeConte, 1847: 83. Type locality: "Pennsylvania." Holotype, 9 , MCZC. Pedilus elegans Hentz; LeConte, 1847: 84. Pedilus newmani LeConte, 1855: 274 (replacement name for lugubris Newman, 9 1838); Abdullah, 1962: 219 ("lectotype" designation...see Type Information, below). Pedilus elegans (Hentz); LeConte, 1855: 274 (synonymy); Long, 1920: 161 (catalog); Abdullah, 1964f: 92-94 (distribution, bionomics). Corphyra elegans (LeConte); LeConte, 1867a: 65 (species list). Corphyra fulvip33(Newman); LeConte, 1867 a: 65 (species list). Corphyra newmmi(LeConte); LeConte, 1867a: 65 (species list). Corphyra fulvipes Newman; Horn, 1871: 281; Horn, 1883: 309, Figure 18; Horn, 1335: 7 (synonymy); Dury, 1902: 177. ' 166 Corphyra newmani LeConte; Horn, 1871: 279; Horn, 1883: 309, Figure 18; Horn, 1885: 7 (synonymy). Corphyra elegans Hentz; Horn, 1871: 282-283; Horn, 1883: 308, Figure 18. Pedilus (Corphyra) elegans Hentz; Pic, 1911: 12 (catalog). Pedilus (Corphyra) fulvipes Newman; Pic, 1911: 12 (catalog). Pedilus (Corphyra) newmani LeConte; Pic, 1911: 13 (catalog). Pedilus fulvipes (Newman); Leng, 1920: 162 (catalog). Pedilus medias (in part) of Abdullah, 1964b: 156, tb. 3, Figures 29-34. Pedilus impressus (in part) of Abdullah, l964f: 91, Figures 13-19. Type Information Two males in LeConte's series of elegans are provided with pink disks, indicative of "Middle States," including Pennsylvania, the type locality listed by Ziegler for haemorrhoidalis. The first specimen agrees most closely with Ziegler's description; it also bears the label, presumably in LeConte's handwriting, "P. elegans (th); 0' haemarrhoidalis ZiegL; 9 ruficollis Ziegl." Presumably, this is the holotype of haemorrhoidalis. A female in the combined Melsheimer - Ziegler collection bears the label, "ruficollis Z. Pa." The abdomen is now missing and certain details, such as the color of the first two antennal segments, do not correspond exactly with Ziegler's very brief description. However, this specimen may possibly be the holotype of ruficollis. LeConte (1855: 274) prOposed P. newmani as a replacement name for P. lugubris Newman, 1838, which is a junior homonym of Anthicus lugubris Say, 1827. Subsequently, Abdullah (1962: 219) designated a specimen from the 167 LeConte collection to serve as the lectotype of the replacement name, newmani LeConte. His action was completely contrary to the rules of zoological nomenclature. In the first place, the type of a replacement nominal Species must be that of the prior nominal species. Furthermore, Abdullah's action was in opposition to the basic definition of a lectotype (i.e., one of the syntypes). On this basis, I consider Abdullah's designation (lectotype, 0' , #30493, MCZC) totally invalid. ‘ The Specimen designated by Abdullah (1964b: 156) to serve as the allotype of his medias (= labiatus) is, in reality, a female of elegans. The Specimen is in the Canadian National Collection (CNCI). Specimens attributed to impressus by Abdullah (1964f: 91) actually belong to elegans. Description Male: Length 5.5-8.5 mm. Head, ventrally exposed portions of mesothorax and metathorax black; prothorax yellowish—orange to reddish-orange, orange suffused to varying degrees with piceous to black, commonly in center of disk, or entirely black; scutellum rufopiceous to black; legs concolorous, testaceous to rufotestaceous or black; abdominal sternites 3-6 piceous to black, 7-8 testaceous to rufotestaceous. Head covered with moderately elongate erect setae, surface finely and sparsely punctulate; mouthparts, labrum and anterior portion of clypeus testaceous to piceous. Antennae weakly serrate, nearly filiform, .articles 1-2 testaceous to rufotestaceous, 3-11 rufopiceous to black. Pronotum ovate, covered with moderately elongate, retrorsely semierect setae, surface finely and Sparsely punctulate. Legs with prothoracic and mesothoracic tarsi conspicuously 168 tomentose beneath; tarsal claws (Figure 42) strongly toothed at base. Elytra moderately coarsely, densely punctate, black with pruinose luster, apices creamy yellow to testaceous; apices impressed, swollen immediately anterad of impressions, impressed and swollen areas impunctate, shining. Abdomen with posterior margins of sternites 7-8 broadly emarginate. Parameres (Figures 83, 117) fused proximally, free and gradually divergent along distal 2/5, each dorsomesally twisted and tapered to a slightly excurved, acutely rounded apex, each bearing a well developed dorsomesally directed subapical tooth from the inner ventral surface; median lobe (Figures 117-118) slightly excavated ventrally, apex forming an ovate knob-like process. Female: As for male except tarsi of prothoracic and mesothoracic legs scarcely more tomentose beneath than those of metathoracic legs; elytra concolorous black with pruinose luster; elytral apices not modified. Abdominal sternites 3-8 concolorous, piceous to black, hind margins of sternites 7-8 entire. Lateral extension of each ventral bacculus (Figure 64) strongly reduced, visible only as a thickening of the bacculus. Distribution (Figure 157) LAURENTIAN: MI, MN, ON, PQ, WI; NEW ENGLAND: CT, MA, ME, NH; MIDDLE ATLANTIC: NJ, NY, PA; MIDDLE STATES: IA, IL, IN, KS, MO, NE, OH; PRAIRIE: MB, ND. [AMNH, BMNH, CASC, CCEC, CDAE, CISC, CNCI, CUCC, CUIC, DBUM, DEFW, DEUN, DYCC, DZEC, EGRC, EJKC, FMNH, GHNC, INHS, JBJC, JLCC, JSCC, KSUC, MCZC, MSUC, NDSU, NMDC, NMPC, OSUC, PADA, PMNH, PSUC, PURC, ROMC, SEMC, UADE, UASM, UCDC, UCRC, UMMZ, UMRM, USNM, WSUCJ Specimens examined: 750 0'0‘ , 166 99 . 169 The interval of time between 15 May and 25 June has been most productive for collecting this species. Diagnosis The weakly serrate, subfiliform antennae, finely and Sparsely punctate head and pronotum, and piceous to black elytra with creamy light yellow to testaceous apices serve to separate males of elegans from all but those of terminaliso The elytral surface of elegans is moderately coarsely, densely punctate and abdominal sternites 7-8 are testaceous to rufotestaceous. Males of terminalis have the elytral surface coarsely and moderately Sparsely plmctate, while abdominal sternites 7-8 are black or occasionally black with testaceous to rufotestaceous apices. Of all the females with a well developed tooth associated with the base of each tarsal claw, only those of elegans have the lateral extension of each ventral bacculus reduced and visible only as a lighfly sclerotized baccular plate. Remark The extreme amount of intraspecific color variation in the pronotum and legs, and sexually dimorphic color differences associated with the elytral apices and abdomen have led to a proliferation of names for this common northeastern species. Varying shades of orange dominate pronotal coloration among the southern and western populations; black is the typical pronotal color of elegans in the extreme northeastern region. . Although Specimens with orange pronotal pigmentation nearly always have light-colored legs, a great deal of variation in leg color may be seen in specimens with a black pronotum. 170 Bionomics Specimens of elegans were reported from hawthorn blossoms in Ontario by Abdullah (1964!). Label data provide several new plant associations as well. At Whitmore Lake in southern Michigan, both males and females were beaten from Cornus racemosa; it has also been collected on Pastinaca sativa in Murray County, Minnesota, on Spiraea flowers at Ames, Iowa, and from flowering Viburnum in lngham County, Michigan. The author has taken it commonly on Crataegus flowers in southern Michigan and by sweeping Bromus inermis. Numerous additional records come from malaise and rotary flight traps. In a brief 1894 note, Harrington alluded to an association between males of elegans, under the name Corphyra newmani LeConte, and meloid beetles: Specimens were twice found mounted upon Meloe niger but for what purpose was not apparent unless they were attracted by the oil exuded by the blister beetle. Males of elegans have been observed at cantharidin bait on numerous occasions in Michigan; additional material, including one female, has been received from Claude Chantal in Quebec. The tiger beetle, Cicindela serguttata(FabriciuS), was observed by the author attacking and feeding upon a male elegans in the vicinity of cantharidin bait in Branch County, Michigan. Approaching from the front, the predator grasped the Pedilus by the head and fed upon the contents of the head and thorax; the elytra and abdomen were discarded. 171 17. Pedilus m SP. NOV. (Figures 34, 153) Type Information Holotype: (0' ), CAL. Burn w. G. [enerall Grant [tree, Grant Grove, Kings Canyori NP, 6-16-64, PSB, P.S. Bartholomew Collection, Calif. Acad. Sci., Accession 1967, [California Academy of Sciencesi. Paratype: (l 0' ), Kings Canyon NP, 6-24-1955, Cal, PS Bartholomew, P.S. Bartholomew Collection, Calif. Acad. Sci., Accession 1967, [CASC]. Description Male: Length 6.5mm. Head black, covered with moderately elongate erect to retrorsely semierect setae, surface moderately finely and sparsely punctate; mouthparts, labrum and anterior margin of clypeus rufotestaceous. Antennae serrate, article 2 and base of article 1 rufotestaceous, remainder of segment 1 black, segments 3-9 rufopiceous, 10-11 rufotestaceous. Prothorax with pronotum piceous, ovate, maximal width 7/5X mesal length, covered with moderately elongate, retrorsely semierect setae, surface moderately finely and sparsely punctate, hypomera rufotestaceous suffused with piceous pigmentation, prosternum black; scutellum and ventrally exposed protions of mesothorax and metathorax black. Legs with coxae and femora black, trochanters, tibiae and tarsi rufotestaceous; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws each bearing a moderately well developed basal tooth. Elytra finely and densely punctate to rugose, rufotestaceous with apices black, each elytron with a 172 yellowish—orange preapical reniform Spot, elytra impressed preapically, swollen around impressed area; impressed and swollen areas impunctate, Shining. Abdomen black, distal margins of sternites 7-8 broadly, shallowly emarginate. Parameres (Figure 84) fused proximally, free and divergent along distal 7/15, each flattened, widely rounded and Slightly excurved apically, each bearing a single dorsoanteriorly projecting spine from the ventrointerior surface; median lobe tapered distally, abruptly expanded at apex, forming a small knob. Female: The female remains to be discovered. Etymology The epithet is a genitive of Grant, referring to the type locality, near the "General Grant" sequoia tree in the Grant Grove region of King's Canyon National Park. Distribution (Figure 158) CALIFORNIAN: CA. See type information for Specific localities. [CASC] Specimens examined: 2 60' . 1'8 . The males of granti possess subapically excurvate parameres, as do those of elegans, lewisi and picipennis- Only the males of granti and lewisi also posses serrate antennae, a well developed tooth associated with the base of each tarsal claw, and preapically impressed elytral apices. The parameres of granti each bear a single, small, dorsoanteriorly projecting spine from the ventrointerior surface (Figure 84); those of 1910131 each bear a Single, well developed, anteromesally curved subapical tooth from the inner margin (Figure 85). 173 Remark Locality data associated with the holotype presumably refer to a small area west of the General Grant sequoia tree which was burned over some years ago. The area was visited during June, 1978, and a small meadow just west of the Grant Grove parking area was discovered. Extensive sweeping and baiting with cantharidin over a two day period yielded no additional Specimens. Bionomics Nothing is presently known regarding the bionomics of granti- Although baiting with cantharidin proved unsuccessful during the visit to the type locality, males probably orient positively toward the chemical as do those of the closely related lewisi- 174 18. Pedlus lewisi (Horn) (Figures 22, 85, 119, 159) Corphyra lewisii Horn, 1871: 281. Type locality: Colorado. Lectotype and paralectotype designated by Abdullah, 1962: 219; lectotype, 0' , 33039, MCZC; paralectotype, 0' , #8178, MCZC. Corphyra lewisi Horn; Horn, 1883: 308, Figure 13. Corphyra lewisi variabilis of Horn, 1883: 308. Type locality and repository unknown (see Type Information, below). NEW SYNONYII. Pedilus (Corphyra) lewisi Horn; Pic, 1911: 13 (catalog). Pedilus arizonensis Fall, 1915: 21-22, Figures 6b, 15; Leng, 1920: 161 (catalog): Abdullah, 1962: 213; Abdullah, l964f: 95-96, Figures 33-33, map 4. Type locality: "Mt. Lemon, Santa Catalina Mts., Arizona." Holotype, 0', #24313, MCZC. NEW SYNONYM. Pedilus lewisi (Horn); Leng, 1920: 162 (catalog); Abdullah, 1962: 219 (designation of lectotype); Abdullah, 1964f: 94-95, Figures 27-32, map 3 (distribution, bionomics). Pedilus lewisi variabilis (Horn); Leng, 1920: 162 (catalog). Type Information The lectotype is from the Horn collection and the paralectotype, referred to as a "paratype" by Abdullah (1962: 219), is from LeConte's series. While this author cannot resolve whether these are actually from Horn's syntype series, it does seem inconsistent that Abdullah did not designate as paralectotypes the remaining lewisi from Colorado in Horn's (1 9 ) and LeConte's (2 O’O' , 2 99 ) 175 collections, particularly since Horn referred to the female in his original description. There are no specimens in either the Horn or LeConte collections labelled as lewisi variabilis; the name resurfaces but one additional time in all subsequent literature (Leng, 1920: 162). From the distribution Horn noted for variabilis, there can be little doubt that he applied the name to males of lewisi which lacked yellowish pigment surrounding the impressed region of the elytral apices. Description Male: Length 5-9 mm. Color of head, scutellum, ventrally exposed portions of mesothorax and metathorax, and abdominal sternites 3-8 piceous to black; pronotum usually yellowish-orange to reddish-orange, infrequently entirely black; prosternum usually yellowish-orange to reddish-orange, commonly with piceous to black hind margin, concolorous black in specimens with black pronotlun. Head covered with moderately elongate erect setae, surface finely and Sparsely punctate. Antennae (Figure 22) serrate, distal portions of articles 1-2 usually testaceous, segments 3-11 rufopiceous to black. Pronotum ovate, covered with moderately elongate, retrorsely semierect setae, surface finely and sparsely punctate. Legs usually rufopiceous to black, sometimes with distal portions of coxae, femora and tibiae more lightly pigmented than remainder; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws strongly toothed at base. Elytra finely and densely punctate to rugulose, usually black, lacking pruinose luster, elytral apices impressed juxtasuturally, swollen around impressed area with swollen region black or yellowish; infrequently elytra concolorous, testaceous. Abdomen with hind margins of 176 sternites 7-8 broadly emarginate. Parameres (Figure 85) fused proximally, free and divergent along distal 4/ 7, each flattened, widely rounded and usually excurved apically, each bearing a Single well developed anteromesally curved subapical tooth from the inner margin; median lobe (Figure 119) with apex tapered to form a bluntly rounded, slightly knobbed point. Female: As for male except tarsi of prothoracic and mesothoracic legs but slightly more tomentose beneath than those of metathoracic legs; elytra usually concolorous black, infrequently entirely testaceous; elytral apices not modified. Abdominal sternites 7-8 with distal margins evenly rounded, entire. Lateral extension of each ventral bacculus well developed. Distribution (Figure 159) SOUTHWEST: AZ, NM; MOUNTAIN: CO, UT, WY; (Doubtful records: CA, GA). [AMNH, BMNH, CASC, CISC, CNCI, CUIC, DCCC, DEFW, DEUN, DYCC, EMUS, FMNH, GHNC, INHS, LACM, MCZC, MSUC, OSUC, PADA, SEMC, SMCC, TAMU, UASM, UCDC, UMRM, USNM.] Specimens examined: 221 0'0' , 50 99 . The peak flight time for lewisi corresponds with that of abnormis : late June to early or middle July. Diagnosis Only the males of 1610131 and granti possess the following set of characters: antennae serrate; tooth of tarsal claw well developed; elytral apices impressed; parameres subapically excurvate. Each paramere of lewisi bears a single, well developed, anteromesally curved subapical tooth from the inner margin (Figure 85); the process associated with each paramere of granti is small and spine-like, 177 dorsoanteriorly projecting, and arises from the ventrointerior surface (Figure 84). Females of lewisi may be distinguished from other Region 2 (see Figure 9) females by the following diagnostic set: antennae rufopiceous to black with distal portions of articles 1-2 usually testaceous; dorsal surface of head and pronotum finely, moderately sparsely punctate; basal tooth of each tarsal claw well developed; sutural angle of each elytron rounded, not dentiform. Remark There can be little doubt after reading Horn's 1883 paper that he confused three species, joanae. lewisi and monticolus. under the specific epithet of lewisi. It is difficult to understand why he was unable to make the distinction between lewisi and his monticolus (= joanae + monticolus) for, as pan pointed out, the male genitalia are considerably different. Having examined the type of Fall's arizonensis, 1 am confident that it is merely a color morph of lewist In addition to the color difference, Fall also stated that arizonensis was smaller, the elytral apices were not appreciably swollen and the impressed area was, "relatively a little smaller." These differences fall well within the intraspecific variability observed in typical specimens of lewist In addition to the pronotal color variability which led to the description of arizonensis, five specimens were examined from Arizona which have testaceous elytra; all previous descriptions mention only black elytra for 19 wisi. The pronotal color is orange in four of these and black in the fifth. The Wyoming specimens represent a new state record. 178 Bionomics A series of Six females and one male were collected by "beating scrub oak" at Durango, Colorado; at Canon, Colorado 3 single male was taken at Pinus ponderosa. The only additional source of general information is provided by a "malaise trap" label attached to a male lewisi from Wayne County, Utah. The author collected three males at cantharidin bait in the Medicine Bow National Forest area of Albany County, Wyoming. The baits were set beneath large Pinus ponderosa a short distance from a small mountain stream. THE LABIATUS GROUP Diagnods Like the lewisi, lugubris and terminalis Species groups, the five species comprising the labiatus group have the parameres gradually and narrowly separated. Unlike the lewisi group, members of the labiatus group have the parameres (Figures 86-90) tapered distally and not at all excurved. Males of the labiatus group have serrate or pectinate antennae, whereas those of lugubris and terminalis group males are nearly filiform. Remark This group is widely distributed in North America; two species, joanae and labiatus have the broadest geographical distributions of any Species in the genus. 179 180 19. Pedilus lauflolsslfall (Figures 47, 86, 120, 160) Pedilus longilobus Fall, 1915: 18-19, Figures 6a, 9; Leng, 1920: 161 (catalog); Abdullah, 1962: 219; Abdullah, l964b: 160 (distribution). Type locality: Plumas Co., California. Holotype, 0' , #24319, MCZC. Description Male: Length 6-8 mm. Head usually concolorOIs black, occasionally with palpi, labrum and anterior portion of clypels testaceOIs, covered with moderately elongate, retrorsely semierect setae, surface finely and Sparsely punctate. Antennae serrate, articles 1-2 rufotestaceous, segments 3-11 piceOIs to black. Pronotum ovate, entirely yellowish-orange to reddish-orange, entirely black, or orange sufflsed with piceOIs or black pigmentation, covered with moderately elongate, retrorsely semierect to decumbent setae, surface finely and sparsely punctate; prosternum entirely yellowish-orange to reddish-orange, entirely black, or orange anteriorly and black posteriorly; scutellum and ventrally exposed portions of mesothorax and metathorax piceOIs to black; legs usually black, infrequently testaceous or testaceOIs suffused with piceous pigmentation; prothoracic and mesothoracic tarsi conspicumsly tomentose beneath; tarsal claws (Figure 47) each with an elongate, apically truncate lobe which is nearly as long as the claw itself. Elytra finely and densely punctate to rugulose, lacking pruinose lister, concolorOIs black or testaceOIs with black apices, sutural margins and occasionally lateral margins; apices flatly impressed, elytra slightly swollen 181 immediately anterad of impressed apices, swollen and impressed areas impunctate, shining; each elytral apex acutely rounded. Venter of abdomen tsually black, infrequently testaceous to piceous, sternites 7-8 with hind margins nearly entire. Parameres (Figure 86) ftsed proximally, free and gradually divergent along distal 1/ 3, each tapering to an acutely rounded apical point, each shallowly notched subapically along the inner margin; median lobe (Figure 120) with apex abruptly narrowed, forming a Slightly knobbed appendix. Female: As for male except tarsi of prothoracic and mesothoracic legs but Slightly more tomentose beneath than those of metathoracic legs; tarsal claws strongly toothed at base; elytra not modified. Lateral extension of each ventral bacculus well developed. Distribution (Figure 160) PACIFIC NORTHWEST: OR, WA; CALIFORNIAN: CA, NV. [BMUW, CASC, CISC, CNCI, DCCC, DYCC, FMNH, JBJC, JSCC, MCZC, MSUC, NHMB, OSUC, OSUO, SMCC, UCDC, UCRC, UMMZ, USNM.] Specimens examined: 85 0'0' , 3 99 . Most of the adults seen were collected between the last week of May and the end of June. Diagnosis The elongate, distally trtmcate tooth of each tarsal claw (Figure 47) is a Specialization unique to the males of longilobus . Females are distinguishable by the following set of characters: antennal articles 1-2 testaceous to rufotestaceOIs; dorsal surface of head finely and moderately sparsely punctate; tooth of tarsal claw well developed; elytral apices rounded, not at all produced. 182 Remark According to Fall, "the male genitalia of [longilobus and serratus are . . . virtually identical." 1 must take exception to this statement, for while the genitalia are Slightly similar between the two, those of longilobus Show a far greater likeness to those of monticolus, labiatus and joanae. Abdullah also recorded longilobus from Alberta. While no Specimens have been examined to confirm the record, its occurance in this province would certainly be expected. Bionomics Plant associations from Corvallis, Oregon, include "sweeping grass" and "flowers of Crataegus douglasi." A dry, rocky, sparsely wooded slope in El Dorado County, California, produced Specimens of longilobus at flowers of Veratrum californicum , a "prostrate white flowering Ceanothus " and an "odd composite." Several males of longilobus have been observed and collected at cantharidin halt in Oregon and California. 183 20. Pedilus crotchi (Horn) (Figures 23-24, 87, 121-122, 161) Corphyra crotchii Horn, 1374: 41. Type locality: Crystal Springs, California. Lectotype, designated by Abdullah, 1962: 213, o' , #3034, MCZC. Corphyra crotchi Horn; Horn, 1883: 307, Figure 16. Pedilus (Corphyra) crotchi Horn; Pic, 1911: 12 (catalog). Pedilus crotchii Horn; Fall, 1915: 15-16, Figures 2, 6c. Pedilus crotchi (Horn); Leng, 1920: 161 (catalog); Abdullah, 1962: 219 (designation of lectotype); Abdullah, 1966b: 181, Figures 43-47. Type Information Abdullah incorrectly listed the number accompanying the lectotype as "#3093" (1962: 219) and later as "no. 3039" (1966b: 181). He also listed a male from the LeConte collection (#7981) as a paralectotype (Abdullah, 1962: 219). All specimens of this species in the LeConte and Horn collections have been examined, there being 3 (l 0' , 2 99 ) in LeConte's series and 4 (l o' , 3 99 ) in Horn's material. The females in Horn's series are labelled "Para - Type; Horn Coll 87879,) but cannot belong to the type series Since Horn clearly stated in the original description that the female was unknown. The male in Horn's series was selected by Abdullah to serve as the lectotype; it also bears the "Horn Coll #7879" label, this leaving this author somewhat doubtful as to whether it actually belongs to the syntype series. Both the lectotype and paralectotype agree in general with Horn's original description; Horn did not mention how many Specimens actually comprised the type series. 184 Description Male: Length 5-8.5 mm. Head black with labrum, distal 1/3 of clypets and frequently first three segments of maxillary palpi yellowish to testaceous, vestiture consisting of short semierect to erect setae, postocular aspect of genae coarsely and moderately densely to densely punctate. Antennae (Figure 23) strongly pectinate beyond 2nd segment, apical portions of articles 1-2 yellowish to testaceOIs, remainder of antennae black. Pronotum ovate, concolorous orange, covered with moderately elongate semierect to erect setae, surface moderately coarsely, sparsely punctate; prosternum orange with posterior margin black; scutellum and ventrally visible portions of mesothorax and metathorax black. Legs with coxae black, remainder variegated, testaceOIs and piceous to black; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws strongly toothed at base. Elytra finely to moderately coarsely, densely punctate, stramineous to testaceOIs with narrow black sutural, apical and lateral margins; apices slightly impressed. Venter of abdomen variegated, piceOIs to black and yellowish to amber, posterior margin of 7th sternite nearly truncate, that of the 8th sternite slightly emarginate. Parameres (Figure 87) ftsed along proximal 3/5, thence divergent and becoming free and subparallel along distal 1/4 of their length, each bearing a well developed, inwardly directed subapical tooth from the inner dorsal surface; median lobe (Figure 121) bearing numerous rows of phallic pectinations (Figure 122) ventromesally, apex gradually tapering to a blunt point. 185 Female: AS for male except antennae (Figure 24) serrate; tarsi of prothoracic and mesothoracic legs but Slightly more tomentose beneath than those of metathoracic legs; elytral apices not impressed. Abdominal sternite 7 with distal margin entire, 8th sternite Short, width at base 4X mesal length, hind margin shallowly emarginate. Lateral extension of each ventral bacculls Short. Distribution (Figure 161) PACIFIC NORTHWEST: OREGON: Curry Co., Thunder Rock Cove; 21-VI- 1967; DYCC (l o' ). CALIFORNIAN: CALIFORNIA: 5 mi. E. Kneeland, Humboldt Co.; 27-V1-1969; UCDC (1 o' ). San Mateo Co., Crystal Springs Res., 9. end San Andreas Lake; 4-v-1973; DYCC (10). San Mateo Co., Crystal Springs Res., 3. end San Andreas Lake; 12-v-1973: DYCC, MSUC (44 dd). San Mateo Co., Crystal Springs Res., 3. end San Andreas Lake; 22-v-1973: DYCC (3 o'o' ). San Mateo Co., Crystal Springs Res., S. end San Andreas Lake; 29-V-l978; DYCC, JBJC (190'). San Mateo Co., Crystal Springs Res., 3. end San Andreas Lake; 20-v-1979; DYCC, JBJC (23 dd , 1 9 ). Specimens examined: 92 dd, 1 9 . Diagnosis The pectinate antennae (Figure 23) serve to differentiate males of crotchi from all other Species of Pedilus. This antennal structure is matched only by the males of flabellatus and parvicollis which have flabellate antennae. The pronotum of crotchi is ovate and widest at the middle; that of flabellatus and parvicollis is widest anteriorly. Females of crotchi and serratus possess strongly serrate antennae and a well developed tooth at the base of each tarsal claw. The femora and ventral abdomen are concolorous rufopiceOIs to black in serratus and variegated testaceous to amber and piceous to black in crotchi. 186 Remark At the onset of this study, crotchi was one of the rarest North American Pedilus, with only three males known. This, the 50 topotypes taken by James and Janice Johnson at cantharidin bait during May, 1978, should be considered a significant discovery. The encroachment of San Francisco might well have eradicated crotchi from the type locality were it not for the fact that a large and important reservoir is located at Crystal Springs. The fenced off area has apparently preserved enough habitat to sustain this pOpulation. Single males from Humboldt Co., California, and Curry Co., Oregon, significantly extend the known geographical range of crotchi; the latter is a new state record. I strongly doubt that the females in the Horn and LeConte series (MCZC) of crotchi are actually attributable to this Species. And, contrary to Abdullah's statement that the antennae are, "subserrate in [thd female," the topotype female in the author's collection possesses strongly senate antennae. Bionomics Over 100 Specimens of crotchi have now been taken from cantharidin bait at the type locality near Crystal Springs (see remarks above). James Johnson has kindly offered his extensive habitat notes of the type locality as well as Specific conditions during the visits of 12 and 22 May, 1978. The area sampled is located along a small road south of San Andreas Lake at 400 feet above sea level. The road is in a ravine and parallels a small stream which serves as an outlet to the lake. A Salix -dominated riparian woodland margins the stream. Trees along the road include Quercus agrifolia, U mbellu- laria californica and Aesculus californica , while Rhus diversaloba is the 187 dominant form of undergrowth. Common flowers duringthe May visits included Scrophularia, Aesculus, Mimulus, Calochortus, Iris, Aquilegia, Brodiae, Sambucus, Heracleum, Sonchus, "Vicia7," "blue scroph.," "small white 'pink'," and "hawkweed." The temperature on 12 May was approximately 80°F, with sunny sides and a light breeze; 43 males were collected. On 22 May, the sides were also sunny but a "moderate wind" prevailed, and the temperature was approximately 65°F; 4 males were collected. Similar habitats were baited in Marin County, south of Alpine Lake, but 'no crotchi were observed. 188 21. Pedilus labiatus (Say) (Figures 14-15, 25, 43, 43, 33, 123, 162) Anthicus labiatus Say, 1827: 247-248. Type locality: "Missouri Territory....Taken on the lower Missouri, near Fort Osage." Holotype, 0'(inferred from original description), [not examined, presumably destroyed]. Lagria (Corphyra) labiata of Say, 1835: 189. Pedilus marginicollis Ziegler, 1844: 47. Type locality: Pennsylvania. Holotype,0', MCZC. Pedilus pulcher LeConte, 1847: 84; LeConte, 1855: 273; Abdullah, 1962: 219. Type locality: Kentucky. Holotype,0', #4377, MCZC. Pedilus labiatus (Say); Leconte, 1855: 273; Leng, 1920: 162 (catalog); Jaques, 1951: 155, Figure 382; Abdullah, l964b: 157-159, tb. 4, Figures 44-50 (distribution, bionomics). Corphyra labiata Say; LeConte, 1867a: 65 (species list); Horn, 1871: 281; Horn, 1883: 309-310, Figure 16; Dury, 1902: 177 (bionomics); Blatchley, 1910: 1331, Figure 531. Corphyra pulchar (LeConte); LeConte, 1867 a: 65 (species list; incorrect subse- quent spelling). Corphyra pulchra LeConte; Horn, 1871: 281 (distribution; incorrect subsequent spelling); Horn, 1883: 309, Figure 16 (distribution); Dury, 1902: 177 (bionomics, synonym of labiata). Pedilus (Corphyra) labiatus Say; Pic, 1911: 13 (catalog). Pedilus (Corphyra) pulcher LeConte; Pic, 1911: 13 (catalog). Pedilus medias Abdullah, l964b: 156, tb. 3, Figures 29-34. Type locality: "Onaga, Pottawatomie County, Kansas." Holotype, d, SEMC. NEW SYNONYM. 189 Type Information Preceding the description of marginicollis, Ziegler stated, "Dr. Melsheimer in litt." Presumably, then, the type came from Melsheimer's material. In the combined Melsheimer-Ziegler collection (MCZC), the lead specimen in the labiatus series is a female which bears a "Melsh" label, denoting that it came from Melsheimer's collection (Hagen, 1884: 196), and a "labiata " label, presum— ably in Ziegler's handwriting. The specimen agrees with Ziegler's description and is probably the holotype. Since the description of marginicollis was apparently provided by Melsheimer, it would seem that the epithet Should be attributed to him as Ziegler may have intended. However, a brief diagnosis provided by Ziegler prefaces the Melsheimer description; since it characterizes the Species as adequately as many descriptions of that time, Ziegler must be credited with authorship. According to the original description of medias, the entire type series was comprised of the holotype (0'), allotype ( 9) and 2 paratypes (both 0" ). The repository of the holotype and one paratype was listed as "UK" (= Snow Entomological Miseum, University of Kansas; SEMC); repository for the other two specimens was not listed. All four Specimens of the type series were examined; since each was either mislabelled or misplaced, they are discussed individually: Holotype: o'(of labiatus; Onaga, Kan.; Crevecoeur. [The type label, presumably written by Abdullah, reads: "PARATYPE o' ." I have added my own label below this, indicating that it should be regarded as the holotype.] Type repository: SEMC. 190 Allotype: 9 (of elegans); Ames, Iowa; May 23, 1928; G.S.W. [Affixed beneath Abdullah's label is a red label bearing the Canadian National Collection number: "AlloTYPE; No. 7963." Beneath this, I have added my own "elegans " determination label.) Repository: CNCI. Paratype "l": 9 (of labiatus); Onaga, Kan.; Crevecoeur. [This is a female, not a male as indicated by Abdullah. It turned up in a loan of material from the British Mlseum, but has now been returned to the proper collection (Hayek, in 11th]. Repository: SEMC. Paratype "2": 0‘ (of labiatus); KAN.; T,B.A. Repository: CDAE. Description Male: Length 4.5-12 mm. Head tsually black with mouthparts, labrum and clypeus yellow to testa- ceous, infrequently entirely yellow to testaceous, covered with moderately elongate erect setae, surface moderately coarsely but rather Sparsely punctate. Antennae (Figure 25) serrate, articles 1-2 reddish-orange, reddish-orange suf- fused with black, or concolorous black, segments 3-11 piceous to black. Prono- tum ovate, disk usually yellowish—orange to reddish-orange with a broad black mesal vitta, rarely concolorous yellowish-orange; hypomera usually yellowish- orange to reddish-orange, sometimes piceous to black; prosternum yellowish- orange to reddish-orange, occasionally with piceous to black hind margin; scutellum usually black, rarely testaceous; ventrally exposed portions of meso— thorax and metathorax piceous to black. Legs testaceous, black, or testaceous 191 sufflsed with black pigmentation, particularly at the distal portions of the coxae, femora and tibiae; prothoracic and mesothoracic tarsi conspicumsly dilated and tomentose beneath; tarsal claws (Figure 43) strongly toothed basally. Elytra moderately coarsely and densely punctate to rugose, black with pruinose lister, each with a glabr01s, subcircular impression near the apex, each with the apex acutely prolonged, this forming a dentiform process. Posterior margin of 8th sternite emarginate. Parameres (Figure 88) flsed proximally, free and slightly divergent along distal 2/ 5, each rounded apically and shallowly notched subapically along inner margin; median lobe (Figure 123) with apex slightly knobbed and keeled ventrally. Female: As for male except tarsi of prothoracic and mesothoracic legs but slightly more dilated and tomentose beneath than those of metathoracic legs; elytra not impressed; dentiform elytral process (Figure 48) usually more pro- nounced; hind margin of 8th abdominal sternite entire. Lateral extension of each ventral bacculus well developed. Distribution (Figure 162) LAURENTIAN: MI, MN, WI; NEW ENGLAND: MA; MIDDLE ATLANTIC: MD, NJ, NY, PA; SOUTHEAST: FL, AR, TN; MIDDLE STATES: IA, IL, IN, KS, KY, MO, NE, OH; PRAIRIE: MT, ND, SD. [AMNH, BMNH, CASC, CCEC, CDAE, CISC, CNCI, CUIC, DBUM, DEFW, DEUN, DYCC, DZEC, EGRC, EJKC, FMNH, GHNC, INHS, JBJC, JSCC, LSUC, MCZC, MSUC, NCSU, NDSU, NMDC, OSUC, OSUO, PADA, PSUC, PURC, SEMC, TAMU, UADE, UASM, UCDC, UCRC, UMMZ, UMRM, USNM, VPIC, WSUCJ Specimens examined: 23306 , 337 99 . Throughout the eastern portion of its range, labiatus is most commonly 192 encountered between late May and the middle of June. In the midwestern states, most specimens were collected in the first two week of June. Diagnosis The males and females of labiatus can easily be recognized by the dentiform prolongation of the sutural angle of each elytron (Figure 48). This condition is approached only by alticolus, a species which is restricted to a small portion of the Californian Sierra-Nevada Mountains; labiatus occurs from Montana to the East Coast. Remark lntraspecific color variation is greatest along the western fringe of the distributional range. The general tendency is for decreasing amounts of black pigmentation in association with the head and pronotum. In the case of the latter, the black medial vitta may be entirely lacking. Abdullah misinterpreted this polymorphism and based his medias on the lighter pigmented morph which is relatively common in Iowa and Kansas. It seems rather remarkable that none of the previous students of Pedilus made any particular note of the unique acuminate-dentiform elytral apices exhibited by labiatus. The character is especially valuable in determining females, since the females of very few Species can be identified on the basis of a Single character. The specimens from Arkansas, Florida, Montana and North Dakota all represent new state records. 193 Bionomics Pedilus labiatus has been collected on the composite, Ambrosia trifida (Dury, 1902), and on "greater horseweed," probably Conyza, and "cup-plant," probably Silphium perfoliatum (Blatchley, 1910). Numerous specimens from Ohio, Michigan and Minnesota have been col- lected at malaise traps; Single Specimens from Tompkins County, New York, and Wolsey, South Dakota, were taken at "Jap. beetle trap #41" and "Light trap," respectively. In Tippecanoe County, Indiana, three females were taken while sweeping nettles. The first record of positive orientation toward cantharidin may be inferred from an observation made by Richard Selander (Abdullah, l964b). He discovered a specimen of labiatus near caged individuals of the meloid, Epicauta fabrici (LeConte), in Illinois. During the course of this study, the author observed and collected males of labiatus at cantharidin bait in Lenawee and Hillsdale counties, Michigan. The most productive habitat for baits at the Lenawee County Site was a moderately shaded grassy lane along the edge of a mixed deciduOIs woodlot, which parallels Bean Creek. Similarly, a grassy path between a wet, densely wooded swamp and a mixed deciduOIs forest produced the labiatus males at cantharidin in Hillsdale County. Larvae presumed to be those of labiatus were collected in Minnesota from "under bark of small branches and logs of Quercus sp., Populus sp. and Celtis Sp. and rotten wood of Quercus Sp. (Ebel, 1954). Ebel (in litt.) indicated that the species association was confirmed by rearing a Single larva. 194 22. Pedlus joules SP. NOV. (Figures 26, 89, 124, 163) Type Information Holotype: (0'), ID: Cassia Co., Howell Canyon, 25 June 1977, A.D. Allen; taken at cantharidin bait [California Academy of Scienced. Allotype: ( 9 ), same data as holotype except, sweeping in vicinity of cantharidin bait [CASC]. Paratopotypes: 10 dd , 7 taken at cantharidin bait, 3 sweeping in vicinity of cantharidin bait [ADAC, 2; CASC, 2; MCZC, 2; DYCC, 4]. Paratypes: 356 0‘0‘ , 23 99 as follows: 2 d'o‘: U.S.A., Minnesota, Clay Co., Buffalo River State Park, Malaise Trap, June 14, 1973; 2 db': IDA [HO]: Willow Flat in Cub. Riv. Canyon, Wasatch Mts., July 4, 1952., B. Malkin; C.N.H.M. 1960, Borys Malkin Coleoptera Colln.; 2 0'0', 2 99: Cypress Hills, Sask [atchewali, 8 VI- 1939, AR. Brook; 3 0’0' , 2 99 : Killdeer Mts., Dunn Co. ND, 23 VI 1965; Collected by L. Grochowski; 3 dd, 15 992: Wawawai, Wn, V-14-1933, LW. Bates; 16 o‘b‘ , 7 99 : UTAH Cache Co, Tony Grove Cany, 26-31 Jul 1975, Knowlton 6: Hanson, Malaise trap 7800'; Id , 2 99 : ALBERTA, CAN., Waterton Lakes N. Pk. Lower Lake, VI-19-1962 4189'; C.W. O'Brien, Collector; ZobiCreston, B.C., l7.V.1946, G. Stace Smith; Ex. G. Stace Smith collection, purchased 1960; 3013': Larkin, B.C., 16.V.l932, G. Stace Smith; Ex. G. Stace Smith Collection, purchased 1960; 7 0'0' : ID: Blaine Co., 1.5 mi N. Sun Vly, 13 April 1977, A.D. Allen; Taken at cantharidin bait; 39 dd: ID: Boise Co., Mile High, 4 June, 1976, G.A. Shook; Taken at cantharidin bait; 126 O'o' : OREGON, Benton Co., Willamette Park [Corvallisi, 24 April 1978, P.J. Johnson; Taken at cantharidin 19S bait; 150 0'0': OR: Yamhill Co., McMinnville, K.M. Fender; Taken at cantharidin bait: l6 (29-IV-1977), 15 (l-V-l978), l8 (16-V-l978), l6 (7-V-l977), 20 (7-V-l978), 26 (8-V-l976), l7 (9-V-l977), 22 (18-V-l978). Paratypes are deposited in the following collections: ADAC, BMNH, CASC, CDAE, CISC, CNCI, DEFW, DYCC, EMUS, FMNH, GSCC, JBJC, JSCC, MCZC, MSUC, NDSU, OSUC, OSUO, SEMC, 0802, and USNM. Description Male: Length 5-8.5 mm. Head black, mouthparts, labrum and anterior portion of clypeus testaceOIs to black, covered with moderately elongate erect and retrorsely semierect setae, surface moderately finely and sparsely punctate. Antennae (Figure 26) serrate, distal portions of articles 1-2 testaceous to reddish-brown, basal portions of segments 1-2 and remaining antennal segments piceous to black. Pronotum ovate, yellowish-orange to reddish—orange, orange sufflsed to varying degrees with piceous to black coloration, or entirely black, covered with moderately elongate, retrorsely semierect setae, surface moderately finely and sparsely punctate; prosternum entirely yellowish-orange to burnt orange, entirely black, or orange with piceOIs to black hind margin; scutellum and ventrally exposed portions of mesothorax and metathorax usually piceous to black, occasionally with testaceOIs to rufotestaceous maculations. Legs testaceOIs to black, com- monly piceOIs to black with variable lighter maculations; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws strongly toothed at base. Elytra moderately coarsely and densely punctate to rugose, black, each elytron with a preapical orange Spot, or testaceous to brown, each 196 with piceous to black sutural and lateral margins and apices as well as a preapical yellowish-orange Spot; apices impressed, elytra somewhat swollen surrounding impressed area; swollen and impressed areas usually impunctate, shining; elytral apices each acutely angulate. Abdominal sternites 3,-8 rufotes- taceOIs, variously sufflsed with piceOIs to black pigmentation, or concolorous piceOIs to black; sternites 7-8 with posterior margins broadly, shallowly emar- ginate, that of the 7th often nearly entire. Parameres (Figure 89) flsed proximally, free and subparallel along distal 5/17, each acutely rounded apically and weakly toothed and notched subapically along inner margin; ventral Sims usually well developed mesally; median lobe (Figure 124) with apex abruptly narrowed and prolonged to form a papilliform process. Female: As for male except head rarely concolorols rufotestaceous; tarsi of prothoracic and mesothoracic legs but slightly more tomentose beneath than those of metathoracic legs; elytral apices lacking preapical yellowish-orange to orange Spot, neither impressed nor swollen; abdominal sternites 7-8 with hind margins entire, 8th lsually retracted within 7th. Lateral extension of each ventral bacculls well developed. Etymology This Species is dedicated with much love and gratitude to my dear wife, Joan. In addition to the support and understanding so often given by a spouse, Joanie also sacrificed numerous "nicer" vacations to accompany me in the field and assist with my research. 197 Distribution (Figure 163) LAURENTIAN: MN; MIDDLE STATES: NE; PRAIRIE: AB, MB, MT, ND, SD, SK; MOUNTAIN: CO, UT, WY; PACIFIC NORTHWEST: BC, ID, OR, WA; CALIFORNIAN: NV. [ADAC, AMNH, BMNH, BMUW, CASC, CDAE, CISC, CNCI, CUIC, DEFW, DYCC, DZEC, EMUS, FMNH, GHNC, GSCC, INHS, JBJC, JSCC, MCZC, MNHS, MSUC, NDSU, NMDC, ODAC, OSUC, OSUO, PURC, SEMC, TAMU, UASM, UBCZ, UCDC, UGCA, UMMZ, UMRM, USNM, WSUCJ Specimens examined: 779 d'O' , 92 99 . Diagmds The development of an excavated region, or ventral sinus, on the underside of the fused parameres is lsually sufficient to differentiate males of ioanae from those of all other species. Other Significant character states include serrate antennae, a well developed tooth associated with the base of each tarsal claw, and elytral apices which are subcircularly impressed with the surrounding surface Slightly elevated or swollen. Females of joanae Share the following character set with those of monticolus: dorsal surface of head moderately coarsely and densely punctate; antennae weakly serrate; basal tooth of each tarsal claw well developed; elytral apices rounded, not at all produced. The two Species are allopatric with monticolus occurring only in the Californian Sierra Mountains and joanae widely distributed from the Pacific Northwest, east to Minnesota, but not recorded from California. Remark Although perhaps the most common species of Pedilus in northwestern North America, joanae has until now masqueraded as its close relative, 198 monticolus. Both exhibit a great deal of intraspecific variation as regards coloration of the pronotum, legs and elytra; characters which have misled previOIs authors to some degree. And, while the two Species appear to be strictly allopatric, their respective geographical ranges come close enough to easily mistake them for a Single distributional pattern. Character discontinuity found in the male genitalia together with the observed allopatry indicate that "monticolus" of previOIs authors is actually a complex of the two species. Fall apparently had some misgivings in this regard, as well. Several Specimens of joanae examined from Fall's series of "monticolus" were furnished with such labels as, "may be true monticola as in LeC. coll.; Typ. ?? monticola Horn; monticola var. or near; compare newmani of col. form of monticola; n. Sp. ? near monticola; near my Plumas Co. exs. of monticola but oedeagls a little diff.," all in Fall's handwriting. Bialomics This species has been collected at malaise traps in Pullman, Washington, and Cache County, Utah; a single male from Whitman County, Washington, Was taken at a malaise trap baited with dry ice. Several males from two Oregon locations bear the label "mech. trap." Single males from Creston, British Columbia, 17 miles west of Calgary, Alberta, Spokane County, Washington, and Wasco County, Oregon, were observed and collected while on Populus trichocarpa, Salix, Quercus, and "alfalfa," respectively. Examples of Pedilus joanae from Saskatchewan carried specimens of the following mite taxa (kindly identified by W. Calvin Welbourn): Erythraeidae, 199 Leptus Sp. larvae; Acaridae, deutonymph; Histiostomidae, deutonymphs (speci- mens in acarology collection, Ohio State University and the author's personal collection). The mites were associated primarily with the ventral abdominal surface of the beetles. Larvae of Leptus are parasitic, whereas nymphs and adults are free-living predators in the soil; acarids and histiostomids are phoretic as deutonym phs, while adults are lsually amociated with wet vegetable matter (Welbourn, in litt. The microhabitat of these mites, together with what little may be inferred regarding the generalized life history of Pedilus, suggest that larvae of joanae are associated with some type of moist, decaying vegetative material on or jtst below the soil surface. The mites may attach to adult joanae soon after the beetles emerge from the pupa, perhaps while they are still teneral and relatively vulnerable to "attack." Leech (1934) commented on three separate associations between what he referred to as Pedilus manticolus (Horn) and Meloe niger Kirby, under the name M. montanus LeConte. The British Columbia locality listed by Leech provides came to believe that the species of Pedilus involved was actually joanae. According to Leech, "In each of the three cases the elytra of the Meloe have actually been eaten away by the Pedilus." Males of joanae have been observed and collected in large numbers at cantharidin bait in Oregon by Kenneth Fender and Paul Johnson, in Idaho by Albert Allen and Gary Shook, and in Saskatchewan by Ronald Hooper. 200 23. Pedilus manticolus (Horn) Figures 90, 164) Corphyra monticola Horn, 1874: 41. Type locality: "Calaveras, California." Holotype, 0' (stated in description), may be in MCZC (see Type Information, below). Corphyra lewisi monticola of Horn, 1883: 308. Pedilus (Corphyra) monticola Horn; Pic, 1911: 13 (catalog). Pedilus monticola Horn; Fall, 1915: 19-21, Figures 6b, 10 (distribution). Pedilus monticola (Horn); Leng, 1920: 161 (catalog). Pedilus monticolus (Horn); Abdullah, 1962: 219; Abdullah, l964b: 160-162, tb. 6, Figures 64-70 (distribution, bionomics). Type Information Fall (1915: 20-21) described in some detail his efforts to trace the holotype of monticolus. This author has retraced his steps and concurs with his decision that the holotype is probably the lead specimen in the LeConte collection (MCZC). This specimen agrees relatively well with Horn's description, even in lacking segments 3-11 of both antennae. It is a male, as Horn stated in the original description, and bears the labels: "Cala," "Type 7984," and "C - monticola Horn." Of the two remaining Specimens in LeConte's monticolus series, one is a male longilobus from California while the other is a female from Arizona, probably that of lewisi. It is, of course, quite possible that the type has disappeared, as Fall pointed out. 201 Description Male: Length 6-9 mm. Head black with palpi, labrum and anterior portion of clypets testaceous to piceous, covered with moderately elongate erect and retrorsely semierect setae, surface moderately coarsely and densely punctate. Antennae weakly serrate, articles 1-2 testaceous, segments 3-11 piceous to black. Pronotum Ovate, entirely yellowish-orange to reddish—orange, or entirely piceOIs to black, covered with moderately elongate, decumbent setae, surface moderately coarsely and densely punctate; prosternum concolorous yellowish-orange to reddish-orange, 4 black or orange anteriorly and black along posterior margin; scutellum, ventrally exposed portions of mesothorax and metathorax piceous to black. Legs piceous to black or black with variable testaceous regions; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws strongly toothed basally. Elytral surface finely and densely punctate to rugulose; each elytron black with a testaceOIs to orange subapical Spot, or testaceous with black apex, sutural and lateral margins; elytral apices acute, flatly, subcircularly impressed; elytra swollen immediately anterad of impressed apices, swollen areas impunctate, Shining. Venter of abdomen piceous to black; posterior margin of 8th sternite emarginate. Parameres (Figure 90) fused proximally, free and gradually diver- gent along distal 2/ 5, each tapering to an acutely rounded point apically, each shallowly notched subapically along inner margin; median lobe tapering to near apex, thence abruptly dilated, forming a short knob. Female: As for male except tarsi of prothoracic and mesothoracic legs but slightly more tomentose beneath than those of metathoracic legs; elytra not modified, concolorous black or testaceous with black apices and occasionally 202 sutural and lateral margins, infrequently entirely testaceous; posterior margin of 8th abdominal sternite entire. Lateral extension of each ventral bacculls well developed. Distribution(Figure164) , CALIFORNIAN: CA, Nv. [CASC, CDAE, CISC, DCCC, DYCC, FMNH, JBJC, LACM, MCZC, MSUC, OSUC, UCDC, UCRC, WSUCJ Specimens examined: 84 0' 0' , 2 99 . The best time to collect monticolus is between the middle of June and the first week of July. Diagnosis Males of monticolus have serrate antennae, a well developed basal tooth associated with each tarsal claw, and elytral apices which are subcircularly impressed with the surrounding elytral surface Slightly swollen. This combina— tion of characters separates them from all but the males of joanae. Unlike monticolus, males of joanae have a well deve10ped ventral Sints associated with the fused region of the parameres. The females of monticolus and joanae may be characterized as follows: dorsal surface of head coarsely and densely punctate; antennae weakly serrate; basal tooth of each tarsal claw well developed; elytral apices rounded, not at all produced. The known geographical range of monticolus includes a portion of the Californian Sierra Mountains from Tulare Co. to northeastern Tehama Co.; joanae is widely distributed from the Pacific North- west, east to Minnesota, but not recorded from California. 203 Remark As understood here, the name monticolus should be applied in a far more restrictive sense than that indicated by Abdullah (l964b). The true monticolus is restricted to the Sierra Mountains of California and the Lake Tahoe region of Nevada. PreViOIs records of "monticolus from the Pacific Northwest, eastward to Minnesota and southward along the Rocky Mountains to central Colorado refer, most likely, to joanae. In his 1883 paper, Horn wrongly synonymized his monticolus (= monticolus 4' joanae) under lewisi. Fall (1915) corrected the error, noting the considerable differences in structure of the male genitalia. lntraspecific color variation is great, involving the pronotum, legs and elytra. Color does not appear to vary clinally, for numerOIs variations are present in a series of 40 males collected by the author and his wife near Fish Camp, in southern Mariposa County, California. Bialiomics Little is known about the habits of monticolus. A single male from Nevada County, California, was collected at a malaise trap in a meadow. Although the author and his wife, together with James and Janice Johnson, observed and collected over 50 males of this species at cantharidin bait in El Dorado and Mariposa counties, not a single specimen was observed on nearby flowers Of various Rosaceae, Umbelliferae and Compositae—all of which are typically productive sources for Pedilus. Montane meadows and stream margins at elevations of 5,000-7,000 feet appear to be the most favorable habitats for this species, as demonstrated by collections from cantharidin bait. 204 There is little doubt that the meloid associations reported for this species by Leech (1934) Should actually be attributed to P. joanae. THE LUGUBRIS GROUP Diagnoab The dorsoventrally flattened parameres (Figures 91-92) and eastern geo- graphical distribution set the two Species forming this small group apart from other North American Pedilus. 205 206 24. Pedilus alarm-is (Say) (Figures 10-13, 16-17, 27, 31-32, 35-41, 53, 55-56, 62-63, 65, 91, 125, 134-141, 165) Anthicus lugubris Say, 1827: 246. Type locality: "Missouri...near Fort Osage." Holotype, 9 (inferred from description), [not examined, presumably destroyed. Anthicus collaris Say, 1827: 246-247. Type locality: "Missouri." Holotype, 9 (inferred from description), [not examined, presumably destroyed . NEW SYNONYM. Pyrochroa ? infumata Hentz, 1830: 257. Type locality: "Massachusetts." Holotype, 9 (inferred from description), [not examined, repository unknowd. Lagria (Corphyra) lugubris of Say, 1835: 189. Lagria (Corphyra) collaris of Say, 1835: 189. Pedilus rufithorar Newman, 1838: 375. Type locality: "N. America." Holotype, 9 (inferred from description), [not examined, BMNH. Pedilus imus Newman, 1838: 375; LeConte, 1847: 84. Type locality: "N. America." Holotype, 9 , BMNH. Pyrochroa inornata Randall, 1838: 23. Type locality: "Maine." Holotype, 9 (inferred from description), [not examined, repository unknown]. Pedilus nigricans Ziegler, 1844: 46. Type locality: "Pennsylvania." Holotype, 9 (inferred from description), may be in MCZC (see Type Information, below). Pedilus infumatus Hentz; LeConte, 1847: 84. 207 Pedilus collaris (Say); L8Conte, 1855: 272-273; Leng, 1920: 162 (catalog); Williams, 1938: 261, Figure 41 (labium, maxillae); Abdullah, l964f: 85-88, Figures 1-6, map 1 (distribution, bionomics). Pedilus lugubris (Say); LeConte, 1355: 273; Leng, 1920: 162 (catalog); Abdullah, l964f: 88-90, Figures 7-12, map 2 (distribution, bionomics). Corphyra lugubris Say; LeConte, 1867a: 65 (species list); Horn, 1871: 280; Horn, 1883: 310, Figure 17; Dury, 1902: 177. Corphyra collar-is Say; LeConte, 1867a: 65 (Species list); Horn, 1871: 282; Horn, 1883: 310, Figure 17. Pedilus (Corphyra) collaris Say; Pic, 1911: 12 (catalog). Pedilus (Corphyra) lugubris Say; Pic, 1911: 13 (catalog). Pedilus lugubris Say; Arnett, 1960-63: 743, Figure 1.83. Type Information Two females in LeConte's series of lugubris (MCZC) bear pink disk, indicative of "Middle States," including Pennsylvania, the type locality listed by Ziegler for his nigricans. Both agree fairly well with the original description of nigricans, the one labelled "lugubris 4" perhaps a bit better. This Specimen may be the holotype of nigricans, in that Ziegler's unique types should still be in the LeConte collection (Hagen, 1884: 195; Arnett, 1961: 128). However, it does not have the customary "Ziegler" label and its authenticity is certainly open to question. 208 Ducription Male: Length 5-8 mm. Head, ventrally exposed portions of mesothorax and metathorax, and elytra black; prothorax yellowish-orange to reddish-orange, orange variably suffused with piceOIs to black pigmentation, or concolorous blach scutellum rufopiceous to black; legs usually black, infrequently entirely testaceous, occasionally piceOIs to black with lighter regions; abdominal stern- ites 3-8 usually piceOIs to black, occasionally with lighter maculae. Head covered with moderately elongate, retrorsely semierect setae, sur- ‘ face finely and sparsely punctulate; mouthparts, labrum and anterior portion of clypeus testaceous to piceOIs. Antennae (Figure 27) weakly serrate, nearly filiform, articles 1-2 testaceous to rufotestaceous, remaining antennal segments piceous to black. Pronotum (Figures 31-32) ovate, covered with moderately elongate, retrorsely semierect setae, surface finely and sparsely punctulate (Figure 35). Legs with prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws strongly toothed at base. Elytral surface moderately coarsely and densely punctate to rugose, with pruinose luster; elytral apices (Figure 53) impressed, impunctate Shining. Abdomen with posterior margin of sternites 7-8 broadly emarginate. Parameres (Figure 91) conspicumsly flattened dorsoventrally, ftsed proximally, free and subparallel along distal 10/13, each somewhat abruptly narrowed to a point apically; median lobe (Figure 125) excavate ventrally, bearing a mesal phallic cord, apex tapered to a bluntly rounded knob. Female: As for male except tarsi of prothoracic and mesothoracic legs but slightly more tom entose beneath than those of metathoracic legs; elytral apices not modified. Abdominal sternites 7-8 with hind margins entire. Lateral extension of each ventral bacculus (Figure 63) well developed. 209 Distribution (Figure 165) MARITIME: NB; LAURENTIAN: MI, MN, ON, PQ, WI; NEW ENGLAND: CT, MA, ME, NH, RI, VT; MIDDLE ATLANTIC: DC, DE, MD, NJ, NY, PA, VA; SOUTHEAST: GA (Leng, 1910), SC; MIDDLE STATES: IA, IL, IN, KS, OH; (Doubtful records: UT, AB). [AMNH, BMNH, CASC, CCEC, CDAE, CISC, CNCI, CUCC, CUIC, DBUM, DEFW, DYCC, EJKC, EMUS, FMNH, GHNC, INHS, JBJC, JSCC, LACM, MCZC, MSUC, NCSU, NDSU, NMDC, NMPC, OSUC, PADA, PMNH, PURC,RDWC, ROMC, SEMC, TAMU, UADE, UASM, UCDC, UCRC, UMMZ, UMRM, USNM, UVCC, WSUCJ Specimens examined: 14470'0‘, 382 99. Although adult lugubris have been collected over a broad Span of time, peak activity appears to be between the end of May and the third week of June. Diagnosis Males of lugubris have weakly serrate, subfiliform antennae and concolor- Ols piceous to black elytra. Males of cyanipennis have a similar antennal structure but the elytra are iridescent indigo blue-green to purple. The elytral apices of both Species are modified but the elytral impression of lugubris is apical and rather broad and flat, while that of cyanipennis is subapical, deeply excavate and reniform. Males of impressus also have subfiliform antennae and concolorOIs piceOIs to black elytra; the elytral impression is subapical and juxtasutural. Females of lugubris may be distinguished from other Region 3 (see Figure 9) females by the following diagnostic set: dorsal surface of head and pronotum finely and Sparsely punctulate; elytra piceous to black; sutural angle of elytron rounded, not dentiform; elytral surface moderately coarsely, densely punctate; lateral extension of each ventral bacculus rod-like. 210 Remark The long established name, collaris, must certainly fall under the synonymy of lugubris. Pronotal color includes not only black (= W13 MW) and yellowish-orange to reddish-orange (= 001le auctarum), but a wide range of intermediates. III fact, several examples have been seen in which the pronotum is orange with a medial black vitta analogOIs to that of labiatus; several were even so labelled! An analysis of pronotal color variation as a function of geographical distribution in Michigan (Table 17) and North America (Table 18) offers weak support to a North-South clinal hypothesis. While specimens in collections clearly Show that the black morph is by far the most common form in such northern regions as Canada, Maine, New Hampshire, Vermont and Michigan, it is not the more prevalent form in Minnesota and Wisconsin as one might anticipate. Here, the red morph appears to be the most common. Specimens examined from Alberta and Utah are probably mislabelled. The Georgia record is that of Leng (1910, under the name COPP’D’M 00110133); additional unconfirmed but probable recorts include Kentucky, Louisiana, Nebraska, North Carolina, North Dakota, South Dakota, Tennessee, West Virginia, Manitoba and Newfoundland (Abdullah, l964f). Bionomics Graenicher (1902, 1907, 1909) listed this Species, under the name Corphyra collarl's, as being associated with flowers of Smilax ecirrhata, S. herbacea, S. hispida, Vagnera (= Smilacina) racemosa, and Senecio aureus in Wisconsin. Working in Indiana, Blatchley (1910) added buckeye, "red haw" (probably a 211 Crataegus) and leaves of the May-apple, Podophyllum peltatum. Observations of anthophiIOIs Coleoptera made by Lovell (1915) at Waldoboro, Maine, over a period of more than 10 years included the following for luwbris: Camus altern- if olia, Veratrum viride and Viburnum lentago. Three additional associations, Barbarea vulgaris in New York, and Iris as well as flowers of Prumls virginiana in Massachtsetts were reported by Abdullah (l964f). Several new plant records may be added to the list. These include alfalfa, "apple leaf" and flowers of PM, Spiraea and Trillium grandiflorum. Habitat labels attached to several males from Hennepin County, Minnesota, indicate that lugubris has been collected in poplar and baswood—maple woods as well as in a tamarack swamp. Males of this species are typically the most common Pedilus at cantharidin bait in Michigan and Quebec; they have also been found in association with the meloid, Meloe mgusticollis (personal observation). On 3 April 1977, two lugubris larvae were collected near the Red Cedar River in East Lansing, Michigan. They were associated with the underside of an . U Imus americana log, adjacent to the soil. The largest specimen was located 10- 15 mm. within the decaying portion of the log; the other larva was discovered as it crawled on the nearby soil surface. The larvae were placed individually in small covered plastic petri dishes along with small amounts of wood and soil from the microhabitat, and kept at room temperatures. The largest of the two larvae pupated on 5 April 1977 and emerged eight days later. The adult, a female, was maintained on a honey-water solution in the lab for 11 days. 212 Table 17. Pedilus lugubris (Say): Analysis of Pronotal Color Nbrphs in Michigan. (B - black pronotum; B/R - black pronotum with reddish.margins; R - red pronotum; RIB - red pronotum with black discal spot) Region 2 B Z B/R Z R. Z R/B Total (-n) Upper Peninsula 100.0 0.0 0.0 0.0 327 Oscoda Co. 100.0 0.0 0.0 0.0 1 Missaukee Co. 100.0 0.0 0.0 0.0 1 Iosco Co. 100.0 0.0 0.0 0.0 1 Lake Co. 100.0 0.0 0.0 0.0 2 Osceola Co. 60.0 20.0 0.0 20.0 5 Clare Co. 100.0 0.0 0.0 0.0 4 Gladwin Co. 100.0 0.0 0.0 0.0 2 Mecosta Co. 100.0 0.0 0.0 0.0 2 Isabella Co. 100.0 0.0 0.0 0.0 1 ‘Midland Co. 100.0 0.0 0.0 0.0 8 Bay Co. 100.0 0.0 0.0 0.0 1 Saginaw Co 98.4 1.6 0.0 0.0 1 Sanilac Co. 100.0 0.0 0.0 0.0 1 Kent Co. 100.0 0.0 0.0 0.0 3 Ionia Co. 100.0 0.0 0.0 0.0 1 Clinton Co. 98.0 0.0 0.0 2.0 51 Shiawassee Co 100.0 0.0 0.0 0.0 31 Lapeer Co. 100.0 0.0 0.0 0.0 5 St. Clair Co. 0.0 100.0 0.0 0.0 1 Eaton Co. 100.0 0.0 0.0 0.0 2 Ingham Co. 97.5 0.5 0.5 1.5 5 Livingston Co. 100.0 0.0 0.0 0.0 3 Oakland Co. 100.0 0.0 0.0 0.0 3 Macomb Co. 100.0 0.0 0.0 0.0 l Kalamazoo Co. 0.0 0.0 0.0 100.0 2 Calhoun Co. 0.0 0.0 20.0 80.0 5 washtenaw Co. 100.0 0.0 0.0 0.0 1 Branch Co. 75.0 0.0 0.0 25.0 4 Lenawee Co. 100.0 0.0 0.0 0.0 2 Total (-n) (B - black pronotum; B/R - Z RIB black pronotum with reddish margins; R - red pronotum; R/B - red pronotum.with black discal spot) Z R 213 2 Elk Pedilus luggbris (Say): Analysis of Pronotal Color Mbrphs ' 2 B in Eastern North America. Table 18. Region 1281 276 47 749 28 184 15 55 43 110 37 16 69 17 33 OoOoOoOOoSOoOoOoOoOogo..0o0ooooooooooo ..... . ... . 1 0003000000000000000000000 .0. .0... 0900930007375005801260030 0005000000000000000000000 0000000002000000300000000 0006050000010000000000000 0...... O 0.107160007222002109840070 090913000 72 620 89 03 1 1.. 1.1.1 1 1 m0 mmmmmmwmmma Emmmnmmmmux 214 25. PedluscymipemisBland (Figures 44, 92, 126, 166) Pedilus cyanipennis Bland, 1864: 254-255. Type locality: "Virginia." [Type not examined, repository unknown]. Corphyra cyanipennis (Bland); LeConte, 1867a: 65 (species list). Corphyra cyanipennis Bland; Horn, 1871: 280; Horn, 1883: 309 (distribution). Pedilus (Corphyra) cyanipennis Bland; Pic, 1911: 12 (catalog). Pedilus cyanipennis (Bland); Leng, 1920: 162 (catalog); Abdullah, 1966b: 178, 180, Figures 40-42 (distribution); Abdullah and Abdullah, 1966: 207-210 (abnormath of male genitalia). Description Male: Length 5.5-7.5 mm. Color of head, pronotum, scutellum and all ventrally visible portions of thorax and abdomen black; elytra superficially iridescent purple, aqua—green and rufopiceOIs tints observable under microscopic examination; maxillary and labial palpi and antennae rufotestaceous. Head sparsely setose, postocular aspect of genae finely and sparsely punctate; antennae weakly serrate, nearly f iliform. Pronotum ovate, covered with fine, retrorsely semierect setae, surface finely and Sparsely punctate. Legs with coxae and femora piceOIs to black, remainder rufotestaceous, prothoracic and mesothoracic tarsi somewhat more tomentose beneath than those of metathoracic legs; tarsal claws (Figure 44) strongly toothed at base. Elytra moderately coarsely and densely punctate, each bearing a Single well developed, subapical and juxtasutural, kidney-shaped impression. Posterior margin of 7th 215 sternite very slighfly emarginate mesally; 8th sternite acutely, deeply emar- ginate. Parameres (Figure 92) conspicuOIsly flattened dorsoventrally, flsed proximally, free and subparallel along distal l/ 2, each tapering to a point apically; median lobe (Figure 126) excavate ventrally and bearing a mesal phallic cord, apex tapering to a blunt point. Female: As for male except distal surface of elytra not modified; posterior margin of 7th sternite rounded. Lateral extension of each ventral bacculls well developed. Distribution (Figure 166) LAURENTIAN: PQ; NEW ENGLAND: MA, NH, VT; MIDDLE ATLANTIC: NY, VA. [AMNH, BMNH, CDAE, CISC, CNCI, CUIC, DSZI, DYCC, MCZC, MSUC, UASM, UMMZ, USNM, UVCCJ Specimens examined: 29 dd, 76 952. The majority of cyanipemlis adults were collected between the second week of June and the first week of July. Diagnosis Both the males and females of cyanipennis differ significantly from all other Species of Pedilus on the basis of elytral and leg coloration. The elytra are iridescent indigo blue-green to purple, the coxae and femora are piceOIs to black and the remainder of each leg is rufotestaceous. Remark This species, which is endemic to the Appalachian Mountains, has the narrowest geographical range of any species of Pedilus inhabiting eastern North 216 America. Three females from Mount Sutton in southern Quebec represent the first records of cyanipennis for Canada. Bionomics Information on the natural history of cyanipennis is essentially nonexistent. One female from the Catskill Mountains of New York bears the simple label, "sweeping." A male from East Dorset, Vermont, was observed "under Mac apple leaf, feeding on mites?" Efforts have not yet been directed toward assessing cantharidin orientation in this species. It seems likely, however, that positive orientation toward cantharidin will eventually be demonstrated for males of cyanipennis. THE TERMINALE GROUP Diagnosis This Species group is defined on the basis of the male genitalia. The median lobe is short and has the Sides strongly convergent distally (Figures 127- 129). Remark Two distinct subgroups may be identified: the canaliculatus subgroup which contains only canaliculatus, and the terminalis subgroup which contains terminalis and impressus. The terminalis group has a northeastern geographical distribution pattern. 217 218 26. Pedilus cauliculatus (LeConte) (Figures 3, 28, 34, 45, 93, 127, 167) Corphyra canaliculata LeConte, 1867: 143; LeConte, 1867a: 67 (Species list); Horn, 1871: 283; Horn, 1883: 308, Figure 11; Dury, 1902: 177-178 (bionomics, color variation). Type locality: "Ohio," Holotype, 9 , #4876, MCZC. Pedilus (Corphyra) canaliculatus LeConte; Pic, 1911: 12 (catalog). Pedilus canaliculatus (LeConte); Leng, 1920: 161 (catalog). Pedilus canaliculatus LeConte; Abdullah, 1962: 219; Abdullah, l964b: 152-153, tb. 2, Figures 11-16 (distribution, bionomics). Pedilus sulcatus Abdullah, 1964b: 153, tb. 2, Figures 17-21. Type locality: "Shelby, Orleans County, New York." Holotype, O' , #5106 (Hoebeke, 1978: 17), CUIC. [2 paratypes examined: 10' (CUIC), 19 (CDAE). NEW SYNONYM. Type Information The holotype of canaliculatus is a teneral female, not a male as indicated by Abdullah (1962: 219). According to the original description of sulcatus, one paratype (0‘) was designated, with the same data as the holotype, from the CUIC material. However, at least two paratypes must have been designated since a female was also examined from "Ohio" (CDAE) which was labelled "paratype," the label being in Abdullah's handwriting. 219 Description Male: Length 4-7 mm. Head lsually concolorous black, infrequently with palpi, labrum and anterior portion of clypets testaceous, covered with Short erect setae, surface isodiametrically microsculptured, hind margin of epicranium bearing a short mesal canaliculus. Antennae (Figure 28) weakly serrate, nearly filiform, apical portions of articles 1-2 testaceOIs to reddish-brown, remainder of antennal segments piceous to black. Pronotum ovate, concolorOIs yellowish—orange to reddish-brown or black, sparsely covered with Short recumbent setae, surface (Figure 34) isodiametrically microsculptured, bearing a longitudinal canaliculus mesally; prosternum of same color as pronotum, often with a narrow black hind margin in light colored specimens; scutellum black; ventrally exposed portions of mesothorax and metathorax black; legs black, prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws (Figure 45) strongly toothed at base. Elytra moderately coarsely, densely punctate, black with slightly thickened creamy light yellow apices, the light color extended obliquely anterad along the lateral margin, dark portions of elytra often with pruinose luster. Venter of abdomen usually black, sternites 7-8 sometimes black with varying amounts of testaceous to piceous coloration, 7 th and 8th sternites broadly emarginate along posterior margins. Parameres (Figure 93) fused along proximal 3/5, narrowly separated for 1/5 their length, thence abruptly diverging postero- laterally and terminating as inwardly curved, apically rounded attenuations; median lobe (Figure 127) with lateral aspects strongly convergent subapically and abruptly divergent apically, forming a broadly rounded bulbous apex which is conspicuously excavated ventrally. 220 Female: As for male except tarsi of prothoracic and mesothoracic legs scarcely more tomentose beneath than those of metathoracic legs; elytral apices not swollen, light pigmentation greatly reduced, restricted to extreme apex. Abdominal sternites 7-8 with apical margins entire, lateral extension of each ventral bacculls well developed. Distribution (Figure 167) LAURENTIAN: MI, PQ; NEW ENGLAND: MA, NH, VT; MIDDLE ATLAN- TIC: NJ, NY, PA, VA; SOUTHEAST: NC; MIDDLE STATES: IN, KY, OH. [AMNH, BMNH, CASC, CCEC, CDAE, CISC, CNCI, CUIC, DBUM, DEFW, DSZI, DYCC, FMNH, GHNC, INHS, JBJC, JSCC, LACM, MSUC, NDSU, NMDC, NMPC, OSUC, OSUO, PADA,! PURC, RDWC, TAMU, UADE, UCDC, UCEC, UCRC, UMMZ, UMRM, USNM, UVCCJ Specimens examined: 341 O'O‘ , 633 99 . The peak flight time of canaliculatus appears to coincide with the flowering. of the rosaceous genera Prunus and Crataegus. Diagnosis The isodiametrical microsculpturing of the dorsal surface of the head and pronotum (Figure 34) is unique to males and females of canaliculatus and thus ,iserves to easily distinguish members of this species from any other Pedilus. Remark Although canaliculatus is rather widely distributed in the Northeast, it is poorly represented in many collections. At first, this seemed very odd as the species is exceedingly abundant in south-central Michigan. It appears, however, 221 that the Species has a very "patchy" type of distribution with large localized populations. Dury (1902) also noted this phenomenon: When LeConte described (2. canaliculata . . . only a single one (type) was known. During 1880, I collected hundreds on the blossoms of "white thorn" and "buckeye," but all were females. lntraspecific variation is most readily associated with the pronotum, where coloration varies from yellowish—orange to black. An analysis of pronotal coloration in large series of Specimens collected at two Michigan localities (Table 19) offers no support to the notion that pronotal cOlor may be correlated with sex. However, the figures presented in Table 20 suggest that a North-South cline may be involved, with the black pronotum becoming more common in the northern demes. No evidence was found to support the recognition of a separate species, suicatus, for specimens with a black pronotum. NumerOIs Specimens have been collected at a single locality in which the pronotum ranged from one extreme to the other with numerous grades between. Abdullah also noted that the median lobe is straight in canaliculatus and "bent over its dorsal surfabe" in sulcatus. The distal portion of the median lobe is quite movable in live specimens; whether it is "bent over" or straight in mounted Specimens appears to be due to both muscle contractions upon death and an artifact of drying. Bionomies Dury (1902) observed canaliculatus in large numbers at flowers of "white thorn" (probably Crataegus) and buckeye near Cincinnati, Ohio; Blatchley (1910) also observed this Species at Crataegus flowers in Indiana. The only additional literature record known to the author is that of Abdullah (l964b). He examined specimens from Ithaca, New York, which were taken at the crucifer, Brassica. 222 A number of plant associations, particularly from the Rosaceae, were noted through the course of this work. Those from label data included Crataegus blossoms (Covington, VA), American plum (Shiawassee County, MI), flowers of Spiraea and "under Mac. apple leaf" (East Dorset, VT), on alfalfa (Lawrence County, PA), and from Taraxacum officinale (Tompkins County, NY). The author has observed and collected this species from variats southern Michigan localities on flowers 01' Ribes, Crataegus, Prunus serotina, P. virginiana, Amelanchl'er canadensis, Pyrus, and Viburnum lentago. On one occasion, hundreds of canaliculatus were observed on m The trees were first examined at around 9:00-9:30 a.m. At this time, individuals were inactive and located primarily within the framework of the inflorescences, away from the individual flowers themselves. Within two hours, the sun was Shining directly on the exposed branches and Specimens of canaliculawg were noted to confine themselves largely to one of two microhabitats and activities. The majority (ca. 7596) were located on the flowers themselves. Although most were covered to varying degrees with pollen, they appeared to be more interested in the nectar. The remaining 2596 were observed: running skittishly back and forth over the twigs near the bases of the inflorescences. Upon closer examination, it was noted that the majority of flower visiting specimens were females while nearly all "twig runners" were males. Observations continued for about an hour, with the idea that the males might be involved in precopulatory behaviors. However, no copulation was noted and very few males ever ventured up to the flowers. At this time, the noted behaviors cannot be accounted for. Males of this species orient readily toward cantharidin bait (Figure 3); the author has collected numerous Specimens in Michigan (Clinton, Shiawassee, 223 Ingham, Livingston, and Lenawee counties), and has received others from N.M. Downie in Tippecanoe County, Indiana. The crab spider, M isumenops asperatus (Hentz), was observed feeding upon trapped specimens of canaliculatus on flowering Ribes sp. and Viburmim lentago in Shiawassee County, Michigan, and on flowering Prunus virginiana in Lenawee County, Michigan. 224 Table 19. Pedilus canaliculatus (LeConte): Analysis "Red" and "Black" Pronotal Morphs in Two Michigan Localities (Site A ca. 60 miles north of Site B). Ratio o H H #R.B R:B 2 Black Site A Shiawassee County Rose Lake Wildlife Experiment Station 99 91:42 a 2.17:1 31.62 O'd' 91:43 =- 2.12:1 32.12 Total 182:85 = 2.14:1 31.8% Site B Lenawee County Ousted State Game Area 99 116:12 = . 9.67:1 9.42 dd 31:2 = 15.50:1 6.1% 10.50:1 8.72 Total 147:14 225 Table 20. Pedilus canaliculatus (LeConte): Analysis of "Red" and "Black" Pronotal Morphs in Northeastern North America. #RzB 2 "Black" Quebec 30:16 34.81 Michigan 380:112 22.8% Indiana 57:9 13.62 Ohio 32:5 13.52 226 27. Pedilus 1mm (Say) (Figures 94, 128, 168) Anthicus impressus Say, 1827: 248. Type locality: Pennsylvania. Holotype,d' (inferred from original description), [not examined, presumably destroyed . Lagria (Corphyra) impressa of Say, 1835: 189. Pedilus impressus (Say); LeConte, 1855: 273; Leng, 1920: 162 (catalog); Abdullah, l964f: 91, Figures 13-19 (misidentification, in part, of elegans). C orphyra impressa Say; LeConte, 1867a: 65 (species list). . Corphyra impressus Say; Horn, 1871: 282 (as synonym of collaris). Corphyra brwmea Blatchley, 1910: 1331-1332. Type locality: Vigo Co., Indiana. Holotype, o‘ , #90, Blatchley Collection (PURC). NEW SYNONYI. Pedilus (CWO) impressus Say; Pic, 1911: 13 (catalog). Pedilus brunneus (Blatchley); Leng, 1920: 162 (catalog). Pedilus W Abdullah, 1964b: 156-157, tb. 4, Figures 35-43- Type locality: Algonquin, Illinois. Holotype, o' , [not examined, repository unknown]. NEW SYNONYI. Type Information Following his description of brunnea , Blatchley noted, "Not an immature form, as first supposed..." Having examined the specimen, I must take exception to this statement. There is no doubt that the type is a teneral adult. In addition to being unable to locate the holotype of parvus, attempts to locate any of the numerous paratypes which Abdullah supposedly designated were also unsuccessful. 227 Description Male: Length 4—6.5 mm. Head piceous to black, covered with short erect setae; surface very finely and sparsely punctulate; palpi, labrum and anterior portion of clypeus rufotest- aceous to piceous. Antennae weakly serrate, nearly filiform, most of article 1 and apex of article 2 testaceous, remainder of antennae piceous to black. Prothorax yellowish-orange to reddish-orange; pronotum transversely ovate, its maximal width 7/5X its mesal length, sparsely covered with short, mostly retrorsely semierect setae, surface very finely and sparsely punctulate; scutel- lum and ventrally exposed portions of mesothorax and metathorax black. Prothoracic coxae sometimes testaceous basally, legs otherwise piceous to black, prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws strongly toothed at base. Elytra coarsely, moderately sparsely punctate, black, frequently with slight pruinose luster; each elytron with a subapical, juxtasutural, ovate impression, impressed area impunctate, shining. Venter of abdomen black with distal margins of sternites 7-8 sometimes testaceous to black, 7 th sternite broadly emarginate along posterior margin, 8th sternite shallowly emarginate along hind margin. Parameres (Figure 94) fused along proximal 1/2, narrowly separated for 1/6 their length, thence diverging slightly, each acutely rounded apically, each bearing a single dorsoanteriorly directed, acute tooth-like process from the inner ventral surface 2/ 3 along their length, where the parameres diverge; median lobe (Figure 128) with sides strongly convergent apically, conspicuously excavated ventrally. Female: As for male except tarsi of prothoracic and mesothoracic legs scarcely more tomentose beneath than those of metathoracic legs; elytral apices 228 not impressed; abdominal sternites 7-8 with distal margins entire. Lateral extension of each ventral bacculus well developed. Distribution (Figure 168) LAURENTIAN: MI, MN, ON, PQ, WI; MIDDLE ATLANTIC: NY, PA; MIDDLE STATES: IN, IL, KS, M0, 0H. [CASC, CNCI, CUIC, DBUM,DEFW, DYCC, EGRC, FMNH, INHS, JBJC, JSCC, MCZC, MSUC, NMDC, NMPC, OSUC, PMNH, PSUC, UCDC, ummz, UMRM, wsvc.) Specimens examined: 248 0'0' , 47 99 . The best time to collect this species appears to be during the last two weeks of May and the first week of June. Diagnosis Males of impressus have weakly serrate, subfiliform antennae and con- colorous piceous to black elytra, as do those of lugubris. The elytral impression is ovate, preapical and juxtasutural in the case of impresws, while a broad, flat apical impression is characteristic for lugubris. Females of impressus and terminalis share the following diagnostic set, as compared to other Region 3 (see Figure 9) females: dorsal surface of head and pronotum finely and sparsely punctulate; elytra piceous to black with surface coarsely and moderately densely punctate; sutural angle of elytron rounded, not dentiform. This author is unable to discriminate between females of impressus and terminalis on the basis of external anatomical characteristics. 229 Remarks The specific epithet, impressus, has been overlooked and improperly applied more than any other name attributed to Pedilus. This is due in part to the fact that neither LeConte nor Horn correctly characterized impressus: perhaps Say's type was lost or destroyed before proper comparisons could be made with material in their collections. The confusion has been perpetuated in recent times by Blatchley (1910) and Abdullah (l964b), who have contributed to the species synonymy. Although frighteningly brief, Say's original description does adequately define the male of impressus. Only two eastern species, impressus and lugubris, have black elytra which are "indented" apically. The "densely and irregularly punctured" elytral surface and specific nature of the impressed area as described by Say adequately differentiate impressus from lugubris. As noted above, Blatchley's brunneus is merely a teneral adult of impres- sus. More difficult to comprehend is Abdullah's association of impressus with elegans, particularly in view of the fact that males of elegans always have the elytra tipped with yellowish pigmentation whereas Say clearly stated that the elytra were black in impressus. At any rate, Abdullah's interpretation left him with specimens (i.e., true impressus) for which he had no name. ThlB, he described pawus, the most recent synonym of impressus. Bionomics Blatchley (1910) observed and collected this species in association with the common plant, Podophyllum peltatum. 230 Single males were collected on "grape" and from flowers of Prunus at State College, Pennsylvania, and Dane County, Wisconsin, respectively. Numerous specimens were also recorded from malaise traps during May at several Minnesota locations, and by "sweeping" in Boone County, Missouri, during late April. The author has taken females of impressus/terminalis at Crataegus flowers in Livingston County, Michigan. The first association between impressus and cantharidin may be inferred from Say's (1827) original description: I found this species early in May, attached to the side of a Meloe' mgusticollis, which was perfectly at rest upon the ground, not appearing to be in the slightest degree incommoded by the weight of its temporary parasite, but seeming rather pleased with its society. Males of impressus have been observed at cantharidin bait on numerous occasions in extreme southern Michigan; additional records were received from Ed Riley in Missouri. In Michigan, only cool, moist riparian and relict bog habitats have produced specimens of this species. 231 23. Pedlus tannindis (Say) (Figures 95, 129, 169) Anthicus terminalis Say, 1827: 247. Type locality: Missouri. Holotype,d (inferred from description), [not examined, presumably destroyed . Lagria (Corphyra) terminalis of Say, 1835: 189. Pedilus guttula Newman, 1838: 375; Leconte, 1847: 84; LeConte, 1855: 273-274. Type locality: "N. America." Holotype, o‘ , BMNH. Corphyra terminalis Say; LeConte, 1867a: 65 (species list); Horn, 1871: 282; Horn, 1883: 308, Figure 12; Dury, 1902: 177-178. Pedilus (Corphyra) terminalis Say; Pic, 1911: 14 (catalog). Pedilus terminalis (Say): Leng, 1920: 161 (catalog); Abdullah, l964b: 154-156, tb. 3, Figures 22-28 (distribution, bionomics). Description Male: Length 4.5-7 mm. Head black with palpi, labrum and anterior portion of clypeus testaceous to black, covered with semierect to erect setae, surface finely and sparsely punctulate; antennae weakly serrate, apical portions of articles 1-2 testaceous to reddish-brown, segments '3-11 piceous to black. Prothorax yellowish-orange to reddish-orange; pronotum ovate to transversely ovate, covered with moderately elongate, retrorsely semierect setae, surface finely and sparsely punctulate: scutellum and ventrally exposed portions of mesothorax and metathorax black. Legs piceous to black, prothoracic and mesothoracic tarsi conspicuously tomen- tose beneath: tarsal claws strongly toothed basally. Elytra coarsely and 232 moderately sparsely punctate, black with creamy yellow to straminious apices, black area commonly with slight pruinose luster; each elytron with a subapical, juxtasutural shallow impression, area immediately surrounding the impression slightly swollen, impressed and swollen areas impunctate, shining. Venter of abdomen black, distal portions of sternites 7-8 occasionally testaceous to rufotestaceous; hind margins of sternites 7-8 broadly, shallowly emarginate. Parameres (Figure 95) fused along proximal 1/2, narrowly separated for 1/6 their length, thence diverging slightly, each bluntly pointed apically and giving rise subapically to a prominent, anteromesally directed tooth-like process from the inner surface; median lobe (Figure 129) with sides strongly convergent apically, conspicuously excavate ventrally. Female: As for male except tarsi of prothoracic and mesothoracic legs scarcely more tomentose beneath than those of metathoracic legs; elytra concolorous black with apices not modified; abdominal sternites 7-8 with posterior margins entire. Lateral extension of each ventral bacculus well developed. Distribution (Figure 169) LAURENTIAN: MI, ON, PQ, WI: NEW ENGLAND: MA; MIDDLE ATLAN- TIC: DC, MD, NY, PA, VA, wv: SOUTHEAST: NC; MIDDLE STATES: IL, IN, KY, Mo, on. [AMNH, BMNH, CASC, CCEC, CDAE, CISC, CNCI, CUIC, DBUM, DCCC, DEFW, DszI, DYCC, FMNH, GHNC, INHS, JBJC, JSCC, MCZC, MSUC, NCSU, NMDC, OSUC, PADA, PSUC, PURC, ROMC, SEMC, UADE, UCDC, UCRC, ULIC, UMMZ, UMRM, USNM, VPIC, WSUCJ Specimens examined: 2080b; 1179?. Over the northern part of) its range, this Species is most active 233 between the middle of May and June. Along the southern distributional fringe, most specimens have been taken in the middle of May. Diagnosis Of the male Pedilus which have weakly serrate, subfiliform antennae, only those of canaliculatus, elegans and terminalis have piceous to black elytra with creamy light yellow to testaceous apices. The dorsal surface of the head and pronotum is isodiametrically microsculptured in canaliculatus and finely, sparsely punctulate in both elegans and terminalis. Males of elegans have a moderately coarsely, densely punctate elytral surface; the elytral surface of terminalis is coarsely and moderately sparsely punctate. Abdominal sternites 7- 8 are testaceous to rufotestaceous in elegans and typically black in terminalis, or black with only the distal margins testaceous to rufotestaceous. Females of terminalis and impressus differ from all other Region 3 (see Figure 9) females by the following set of characters: dorsal surface of head and pronotum finely and sparsely punctate: elytra piceous to black with surface coarsely and moderately densely punctate; sutural angle of elytron rounded, not dentiform. A reliable methdd has not been discovered to distinguish females of terminalis from those of impressus. Remarks Abdullah (1964b: 157) contended that the basal tooth of the tarsal claw was "shorter" in females of impressus, but this character is correlated more closely to overall size, with much overlap between the two species. 234 Abdullah (l964b) also listed terminalis from New Jersey and Tennessee. While no specimens were seen to confirm these records, there can be little doubt that it occurs there and in all other states to the north and east. Bionomics Robertson (1892, 1894) reported observing terminalis on flowers of Isopyrum biternatum and Ranunculus septentrionalis, Abdullah (l964b) listed it from Barbarea vulgaris in New York and Camus in North Carolina, and Grant and Grant (1965) collected specimens at flowers of Polemonium reptans. A male from Washington County, Maryland, was collected at "dogwood blossoms;" Similar observations come from Michigan (beating Cor-ms racemosa) and North Carolina ("Camus"). Other Michigan associations include Crataegus and wild Hydrangea blossoms, while a male and female were taken on leaves of Alliaria petiolata in Morgan County, West Virginia. Pinto and Selander (1970) recorded the following associations between terminalis and meloid beetles of the genus Meloe: We have seen a male M. angusticollis from Fayette County, Pennsylvania, and a female M. americanus from Rankin, Missouri, each with an adult of Pedilus terminalis mounted on the dorsum of the body; . . . the Meloe elytra had been chewed (along the sutural margin)." The author has collected males of terminalis at cantharidin bait on several occasions in Livingston County, Michigan, where females of impresws/terminalis have also been observed at flowers of Crataegus. UNPLACED SPECIB The positiom of P. serratus and P. alticolus relative to the six species groups of North American Pedilus remain unresolved at present. In terms of general structure and geographical distribution, both would appear to be most closely related to the punctulatus group. 235 236 29. Pedlus cavatus Fall Pedilus serratus Fall, 1915: 19, Figures 3, 6b, 9a, 9b; Leng 1920: 161 (catalog): Abdullah, 1962: 219; Abdullah, l964b: 159, tb. 5, Figures 51-56 (distribu- tion). Type locality: El Dorado Co., California. Holotype, o' , #24323, MCZC. Description Male: Length 6.5-7.5 mm. Color of head, scutellum, ventrally exposed portions of mesothorax and metathorax and abdominal sternites 3-8 black; ‘ mouthparts testaceous to black; pronotum yellowish-orange or black; prosternum yellowish-orange with black hind margin or entirely black. Head covered with moderately elongate, retrorsely semierect setae, sur- face finely and sparsely punctate. Antennae (Figure 29) strongly serrate, articles 1-2 entirely testaceous; infrequently dark basally and testaceous apical- 1y, segments 3-ll rufopiceous to black. Pronotum ovate, covered with moder- ately elongate, retrorsely semierect setae, surface finely and sparsely punctate. Legs commonly with coxae and femora rufopiceous to black, remaining segments testaceous, occasionally legs concolorous black; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws strongly toothed at base. Elytra moderately coarsely and densely punctate to rugose, testaceous with black apices or concolorous black, lacking pruinose luster, tapering distally; elytral apices weakly, flatly impressed, slightly swollen immediately anterad of impressed area; impressed and swollen regions impunctate, shining. Hind margin 237 of 7th abdominal sternite entire, that of 8th sternite emarginate. Parameres (Figure 96) fused proximally, free and subparallel along distal 5/19, each acutely rounded apically and bearing a single, well developed, anteromesally directed subapical tooth from inner surface; median lobe (Figure 130) bearing numerous rows of phallic pectinatiom ventrally (Figure 131), sides abruptly tapering. distally, apex acutely rounded, slightly knobbed. Female: As for male except antennae (Figure 30) less strongly serrate; tarsi of prothoracic and mesothoracic legs but slightly more tomentose beneath than those of metathoracic legs; elytral apices not modified, neither apically impressed nor swollen subapically. Eighth abdominal sternite retracted within 7th abdominal segment, poorly sclerotized; lateral extension of each ventral bacculus well developed. Distribution (Figure 170) CALIFORNIAN: CALIFORNIA: Towie, Acc. 38903; 25-VI-l933; AMNH (10). Feather River, Carabou; l-VI-l946; CISC (l o' , l 9 ). Chalk Bluff Rd., Nevada Co.; 20-VI-l960; CDAE (2 0' 0' ). White Cloud G.S., Nevada Co.; 20-VI- 1960; CDAE (1 9 ). Carrville, Trinity Co., Alt. 2400-2500 ft., Chicago N.H. Mts.; lG-VI-l934; FMNH (l d ). El Dorado Co.; ll-VI-l906; MCZC (l d ). El Dorado Co.; 12-V1-l9l3; MCZC (10'). El Dorado Co., El Dorado Nat. For., Bridalveil Picnic Gd.; 24-VI-l977; DYCC (1 0' ). Specimens examined: 8 0'0' , 2 99 . Diagnosis The following set of characters will serve to characterize males of serratus: strongly serrate antennae (Figure 29); a well developed basal tooth 238 associated with each tarsal claw; elytral apices weakly, flatly impressed and Slightly swollen anterad of impressed area; parameres tapering, acutely rounded apically, each bearing an acute subapical tooth. Females share the following diagnostic set with those of crotchi: antennae (Figure 30) serrate; dorsal surface of head moderately coarsely and densely punctate; basal tooth of tarsal claw well developed; elytral apices rounded, not at all produced. The femora and abdominal venter of serratus are concolorous, rufopiceous to blaCk; these regions are variegated, testaceous to amber and piceous to black, in crotchi. Remarks Although very few specimens of this uncommon species were available for study, a great deal of intraspecific variation was noted relative to color of the pronotum, legs and elytra. According to Abdullah (l964b: 159), "In the male, the genitalia are similar to that in P. longilobus." I strongly disagree; the male genitalia of serratus are far more suggestive of dentatus while those of longilobus come closest to labiatus and monticolus. This species appears to be restricted to the central and northern portion of the Sierra Mountains and the mountainous regions of northwestern California; Abdullah's Alameda Co., California, record is almost certainly in error. Bionomics Two males and a single female of serratus from the California Department of Agriculture collection were taken at Frick traps in Nevada County, California, on 20 June 1960. According to Kenneth Frick (personal communica- 239 tion), the devices were used extensively by the California Department of Agriculture in the late 1950‘s and early 1960's to survey populations of cherry fruit flies. The Frick trap, actually a modification of Hodson's (1948) design, makes use of a cylinder or board treated with a sticky material in association with ammonium carbonate, an "attractant" (Frick, 1952; Prick, et aL, 1954). The fact that no other Pedilus bear the "Frick trap" label indicates that the association was probably accidental. Field studies with ammonium carbonate have not yet been attempted, but will be investigated in the future. In El Dorado County, California, two serratus males were observed at cantharidin bait. The bait was placed in a grassy area beneath large Acer grandifolia, approximately 10-15 yards from a stream (James Johnson, in nu). 240 30. Pedilus dticolus 12.11 (Figures 49, 97, 132, 171) Pedilus alticola Fall, 1915: 27, Figures 4, 13: Leng, 1920: 161 (catalog). Type locality: "Atwood's Mills, Tulare Co., 6600 feet." Holotype, o‘ , #24312, MCZC. Pedilus alticolus Fall; Abdullah, 1962: 218. Pedilus alticolus (Fall); Abdullah, 1966b: 177-178, Figures 31-36 (distribution). Type Information When the . holotype was examined in September of 1978, the head and prothorax were missing, the damage presumably having been sustained between 1966 and 1978 since Abdullah made no mention of it in his work. Description Male: Length 5.5-9 mm. Color of head, ventrally exposed portions of metathorax, legs, elytra and abdominal sternites 2-8 piceous to black; labrum testaceous to piceous; prothorax concolorous reddish-orange; ventral portions of mesothorax piceous to black, mesosternum partially yellowish-orange in some specimens; sclerotized portions of abdominal segments 9-10 and genitalia yellow- ish-brown to piceous. Head covered with moderately elongate erect setae, postocular aspect of genae finely and sparsely to moderately densely punctate. Antennae serrate, distal portions of articles l-2 usually more lightly pigmented than remainder. Pronotum ovate, covered with moderately elongate retrorsely semierect setae, 241 finely and sparsely punctate; prothoracic and mesothoracic tarsi conspicuously tomentose beneath; tarsal claws strongly toothed at base. Elytral surface finely and densely to rugulosely punctate, lacking pruinose luster; elytral apices each with a single well developed juxtasutural impression which is slightly more than twice as long as its maximum width; elytra bluntly subacuminate apically. Abdomen with 7th sternite broadly and shallowly emarginate along distal margin; distal margin of 8th sternite mesally emarginate. Parameres (Figure 97) fused proximally, free and subparallel along distal 3/8, each tapered to a blunt point apically, each bearing a preapical tuft of anteromesally directed spinulae along the inner margin; median lobe (Figure 132) with apex abruptly narrowed, bluntly rounded. Female: As for male except tarsi of prothoracic and mesothoracic legs but slightly more tomentose beneath than those of metathoracic legs; elytral apices (Figure 49) not modified; elytra bluntly pointed apically. Abdominal sternites 7- 8 with hind margins entire, that of 8th sternite slightly produced posterad at meson; lateral extension of each ventral bacculus well developed. Distribution (Figure 171) CALIFORNIAN: CALIFORNIA: Yosemite Valley; 22-V-l908; CASC (10'). Kings Canyon NP; 24-VI—l955; CASC (2 o'o' ). Tulare Co., 8. Cal., Cornell U. Lot 908, Sub.; [no date]; CUIC (l 0' , 19 ). Kaweah, Homotype compared by Frost 1944; 6-VI; MCZC (l 0' ). Kaweah, Homotype compared by Frost 1944, 6000 ft.; 6-VI; MCZC (l o' ). Round M'd'w, Giant Forest; "July"; MCZC (l 0' ). Kaweah; [no datd; UMRM (l 0'). Sierra N.F.; [no datd; USNM (10' ). Round M'd'w, Giant Forest, Tulare Co.; [no date]; MCZC (19 ). [California, locality not legibld, Ac. 26824; 23-v-1919: AMNH (10'). Specimens examined: 1060', 2 99 . 242 Diagnosis The elongate, narrow juxtasutural elytral impression is diagnostic for males of this rare species. Also unique to males of alticolus is the A tuft of spinulae associated with the inner margin of each paramere. Female alticolus differ from those of other Region 1 (see Figure 9) species by their acutely produced elytral apices; they also have a well developed tooth associated with the base of each tarsal claw. Remarlo The specimens examined show little variability, but the small series available for this study precludes offering any generalizations. As alluded to under "Intrageneric Phylogenetic Considerations (p. 52)," the relationships between alticolus and other species of Pedilus remain poorly understood. The presence of apparent sym plesiomorphies and at least one autapomorphy do nothing to clarify the evolutionary history of alticolus. Bionomics The only information available regarding the natural history of alticolus came from a label reading "wild cherry" which is attached to a male in the collection of the American Museum of Natural History. INCERTAE SEDIS Corphyra calypso Wickham (Fossil species) Corphyra calypso Wickham, 1914: 491, plate 16, Figures 3-4. Type locality: Florissant, Colorado (Miocene deposits). Holotype, sex undeterminable, #2,696, MCZC. Pedilus calypso (Wickham), Leng, 1920: 354 (catalog). Remarks The holotype was examined in as much detail as was possible, but Wickham's conclusions as regards generic position or familial affinities could be neither supported nor refuted. It seems at best conjectural to refer it to Pedilus, but I am unable to find evidence in support of placing it elsewhere. 243 Figure 10. Figure 11. Figure 12. Figure 13. Figure 14. Figure 15. Figure 16. Figure 17. 244 PLATE 9 Pedilus lugubris (Sail): cranium, adult male (dorsal view). Pedilus lugubris (Say): cranium, adult male (ventral view). Pedilus lugubris (Say): cranium, adult male (lateral view). Pedilus lugubris (Say): cranial endoskeleton - tentorium. Pedilus labiatus (Say): right mandible, adult female (dorsal view). pedizus labiatus (Say): right mandible, adult female (ventral view). Pedilus lugubris (Say): right maxilla, adult male (ventral view). Pedilus lugubris (Say): labium, adult male (ventral view). 245 246 PLATEIO Figures 18-24. Antennae of adult Pedilus spp. Figure 18. Pedilus flabellatus (Horn): male. Figure 19. Pedilus parvicollis Fall: male. Figure 20. Pedilus cavatus Fall: male. Figure 21. Pedilus inconspicws (Horn): male. Figure 22. Pedilus lGWi-Si (Horn): male. Figure 23. Pedilus CTOtChi (Horn): male. Figure 24. Pedilus crotchi (Horn): female. Figure 25. Figure 26. Figure 27. Figure 28. Figure 29. Figure 30. 248 PLATE 11 Figures 25-30. Antennae of adult Pedilusspp. Pedilus labiatus (Say): female/ male. Pedilus joanae sp. nov.: female/ male. Pedilus lugubris (Say): male. Pedilus canaliculatus (LeConte): female/ male. Pedilus serratus Fall: male. Pedilus serratus Fall: female. 250 PLATEIZ Figure 31. Pedilus lugubris (Say): pronotum, adult female/male (dorsal view). Figure 32. Pedilus lugubris (Say): pronotum, adult female/male (ventral view). Figure 3:. Pedilus flabellatus (Horn), P. parvicollis Fall: pronotum, adult female/male (dorsal view). Figure 34. Pedilus canaliculatus (LeConte): pronotal microsculpturing, adult female/male (dorsal view). Figure 35. Pedilus lugubris (Say): pronotal microsculpturing, adult female/male (dorsal view). Figure 36. Pedilus lugubris (Say): mesonotum, adult female/male (dorsal view). Figure 37. Pedilus lugubris (Say): right mesothoracic wing. Figure 38. Pedilus lugubris (Say): metendosternite. 251 mesothoracic . . radial cel scutum 36 mesothoracnc scutellum 3 7 subcubital "90k wedge cell Figure 39. Figure 40. Figure 41. Figure 42. Figure 43. Figure 44. Figure 45. Figure 46. Figure 47. 252 PLATE l3 Pedilus lugubris (Say): right prothoracic leg, adult male. Pedilus lugubris (Say): right mesothoracic leg, adult male. Pedilus lugubris (Say): right metathoracic leg, adult male. Pedilus elegans (Hentz): tarsal claws, adult female/male. Pedilus labiatus (Say): tarsal claws, ath female/ male. Pedilus cyanipennis Bland: tarsal claws, adult male. Pedilus canaliculatus (LeConte): tarsal claws, adult female/male. Pedilus inconspicuus (Horn): tarsal claw, adult female/male. Pedilus longilows Fall: tarsal claws, adult male. 253 Figure 48. Figure 49. Figure 50. Figure 51. Figure 52. Figure 53. Figure 54. 254 PLATE l4 Pedilus labiatus (Say): right elytral apex, adult female. Pedilus alticolus Fall: right elytral apex, adult female. Pedilus bardi (Horn): right elytral apex, adult female. Pedilus vittatus (Horn): right elytral apex, adult female. Pedilus punctulatus LeConte: elytral apex, adult male. Pedilus lugubris (Say): surface of elytral organ, adult male. Pedilus cavatus Fall: surface of elytral organ, adult male. 5 255 256' PLATE15 Figure 55. Pedilus lugubris (Say): abdominal tergite eight, adult female. Figure 56. Pedilus lugubris (Say): abdominal sternite eight, adult female. Figure 57. Pedilus rusticus Abdullah: abdominal sternite seven, adult female. Figure 58. Pedilus flexiventris Fall: abdominal sternite seven, adult female. Figure 59. Pedilus johnsonorum sp. nov.: abdominal sternite seven, adult female. Figure 60. Pedilus inconspicuus (Horn): abdominal sternite seven, adult female. Figure 61. Pedilus picipennis Fall: abdominal sternite seven, adult female. Figure 62. Pedilus lugubris (Say): distal ovipositor (dorsal view). Figure 63. Pedilus lugubris (Say): distal ovipositor (ventral view). Figure 64. Pedilus elegans (Hentz), P. vittatus (Horn): apex of ovipositor (ventral view). 257 258 PLATE 16 Figure 65. Pedilus lugubris (Say): abdominal segments 9-10, adult male (dorsal view). Figures 66-81. Parameres of adult male Pedilus spp. (dorsal view). Figure 66. Pedilus tibialis Semenov. Figure 67. Pedilus pallidipennis Semenov. Figure 68. Pedilus flabellatus (Horn). Figure 69. Pedilus parvicollis Fall. Figure 70. Pedilus abnormis (Horn). Figure 71. Pedilus dentatus Abdullah. Figure 72. Pedilus rusticus Abdullah. Figure 73. Pedilus fenderi sp. nov. Figure 74. Pedilus flexiventris Fall. Figure 75. Pedilus cavatus Fall. Figure 76. Pedilus johnsonorum sp. nov. Figure 77. Pedilus inconspicuus (Horn). Figure 78. Pedilus vittatus (Horn). Figure 79. Pedilus oregonus Fall. Figure 80. Pedilus bardi (Horn). Figure 81. Pedilus punctulatus LeCo'nte. 259 Figures 82-97. Parameres of adult male Figure 82. Figure 83. Figure 84. Figure 85. Figure 86. Figure 87. Figure 88. Figure 89. Figure 90. Figure 91. Figure 92. Figure 93. Figure 94. Figure 95. Figure 96. Figure 97. 260 Pedilus picipennis Fall. Pedilus elegans (Hentz). Pedilus granti sp. nov. Pedilus lewisi (Horn). Pedilus longilobus Fall. Pedilus crotchi (Horn). Pedilus labiatus (Say). Pedilus joanae Sp. nov. PLATE l7 Pedilus monticolus (Horn). Pedilus lugubris (Say). Pedilus cyanipennis Bland. Pedilus canaliculatus (LeConte). Pedilus impressus (Say). Pedilus terminalis (Say). Pedilus serratus Fall. Pedilus alticolus Fall. Pedilus spp. (dorsal view). 261 .vfifiem 1 . N.W.... e»... L., .2433... 19.5 262 PLATE 18 Figures 98-102, 104-107. Median Lobe of adult male Pedilus spp. (ventral view). Figure 98- Pedilus flabellatus (Horn). Figure 99- Pedilus parvicollis F911: Figure 100- Pedilus abnormis (Horn). Figure 101- Pedilus dentatus Abdullah. Figure 102. pedflus rusticus Abdullah. Figure 103. Pedilus fenderi sp. nov.: genitalia, adult male (ventral view). Figure 104. Pedilus fenderi sp- DOV- Figure 105- Pedilus flexiventris F811- Figure 106. Pedilus cavatus Fall. Figure 107. Pedilus johnsonorum sp. nov. Figure 108. Pedilus johnsonorum sp. nov.: apex of parameres, adult male (ventral view). 263 Figure 109. Figure 110. Figure 111. Figure 112. Figure 113. Figure 114. Figure 115. Figure 116. Figure 117. Figure 118. Figure 119. 264 PLATE 19 Figures 111-114, 116, 118-119. Median lobe of adult male Pedilus spp. (ventral view). Pedilus inconspicuus (Horn): apex of median lobe, adult male (ventral view). Pedilus inconspicuus (Horn): apex of right paramere, adult male (ventral view). Pedilus vittatus (Horn). Pedilus oregonus Fall. Pedilus bardi (Horn). Pedilus punctulatus LeConte. Pedilus punctulatus LeConte: apex of median lobe, adult male (ventral view). Pedilus picipennis Fall. Pedilus elegans (Hentz): genitalia, adult male (ventral view). Pedilus elegans (Hentz). Pedilus lewisi (Horn). 265 266 PLATE 20 Figures 120-121, 124-125, 127-130, 132. Median lobe of adult male Figure 120. Figure 121. Figure 122. Figure 123. Figure 124. Figure 125. Figure 126. Figure 127. Figure 128. Figure 129. Figure 130. Figure 131. Figure 132. Pedilus spp. (ventral view). Pedilus longilobus Fall. Pedilus crotchi (Horn). Pedilus crotchi (Horn): meson of median lobe, adult male (ventral view). Pedilus labiatus (Say): apex of median lobe, adult male (ventral view). Pedilus joanae sp. nov. Pedilus lugubris (Say). Pedilus cyanipennis Bland: apex of median lobe, adult male (ventral view). Pedilus canaliculatus (LeConte). Pedilus impressus (Say). Pedilus terminalis (Say). Pedilus serratus Fall. Pedilus serratus Fall: meson of median lobe, adult male (ventral view). Pedilus alticolus Fall. 267 268 PLATEZI Figure 133. Pedilus sp., larva (dorsal view). 269 \ \,, I Figure 133. 270 PLATE 22 Figures 134-141. Anatomical features of Pedilus lugubris (Say), larva. Figure 134. Habitus (lateral view). Figure 135. Mesothoracic spiracle. Figure 136. Abdominal spiracle. Figure 137. Epipharynx. Figure 138. Left mandible (dorsal view). Figure 139. Mandibular armature (dorsal view). Figure 140. Right maxilla (ventral view). Figure 141. Labium (adoral surface). 271 272 PLATE 23 Figures 142-144. Geographical Distribution Maps. Figure 142- Pedilus flabellatus (Horn). Figure 143- Pedilus parvicollis F811- Figure 144- Pedilus abnormis (Horn)- 273 274 PLATE 24 Figures 145-147. Geographical Distribution Maps Figure 145. Pedilus dentatus Abdullah. Figure 146. Pedilus rusticus Abdullah. Figure 147. Pedilus fenderi sp. nov. 275 276 PLATE 25 Figures 148-151. Geographical Distribution Maps. Figure 148. Pedilus flexiventris Fall. Figure 149. Pedilus cavatus Fall. Figure 150. Pedilus johnsonorum Sp. nov. Figure 151. Pedilus inconspicuus (Horn). 277 278 PLATE 26 Figures 152-155. Geographical Distribution Maps. Figure 152. Pedilus vittatus (Horn). Figure 153. Pedilus oregonus Fall. Figure 154. Pedilus bardi (Horn). Figure 155. Pedilus punctulatus LeConte. 279 280 PLATE 27 Figures 156-159. Geographical Distribution Maps. Figure 156. Pedilus picipennis Fall. Figure 157. Pedilus elegans (Hentz). Figure 158. Pedilus granti sp. nov. Figure 159. Pedilus lewisi (Horn). 281 Figure 160. Figure 161. Figure 162. Figure 163. Figure 164. Figure 165. Figure 166. 282 PLATE 28 Figures 160-166. Geographical Distribution Maps. Pedilus longilobus Fall. Pedilus crotchi (Horn). Pedilus labiatus (Say). Pedilus joanae sp. nov. Pedilus monticolus (Horn). Pedilus lugubris (Say). Pedilus cyanipennis Bland. 283 284 PLATE 29 Figures 167-171. Geographical Distribution Maps. Figure 167. Pedilus canaliculatus(LeConte). Figure 168. Pedilus impressus (Say). Figure 169. Pedilus terminalis (Say). Figure 170. Pedilus serratus Fall. Figure 171. Pedilus alticolus Fall. 285 BIBLIOGRAPHY BIBLIOGRAPHY Abdullah, M. 1960. A revision of the genus Pergetus (Coleoptera: Pedilidae). Can. J. 2001. 38:875-878, 6 figs. . 1961a. A revision of the gems Leptoremus (Coleoptera: Pedilidae). Ann. Ent. Soc. Amer. 54:73-75, 9 figs. . 1961b. A revisional study of some Australian species of Egestria (Pedilidae). Coleop. Bull. 15:27-30, 13 figs. . 1961c. Systematics of 0111701311”, a new genus of pedilid beetles (Pedilidae). 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Cantharidin orientation in insects: Historical review and recent observations. (in preparation) Ziegler, D. 1844. Descriptiom of new North American Coleoptera. Proc. Acad. Nat. Sci. Philadelphia 2:43-53. APPENDIX APPENDIX: PLANT W110“ FOR NORTH AMERICAN SPECIE OF PEDILUS ADULTS The following list was compiled from all known literature records as well as data extracted from specimen labels and observations made by the author. Specific details of the associations, where known, are reported under the "Bionomics" section for each species. Listing of plant families follows Cronquist (1968). PINACEAE Pinus attenuata Lemmon bardi P. jeffreyi Murray abnormis P. monticola Douglas flabellatus P. ponderosa Douglas inconspicuus, lewisi "pine" punctulatus LAURACEAE U mbellularia sp. cavatus RANUNCULACEAE Isopyrum biternatum (Rafinesque) terminalis Ranunculus septentrionalis Poiret terminalis Ranunculus sp. inconspicuus BERBERIDACEAE Podophyllum peltatum Linnaeus lugubris, impressus URTICACEAE U rtica dioica Linnaeus vittatus "nettles" labiatus FAGACEAE Quercus ogrifolia Nees Q. lobata Nees Quercus Sp. "scrub oak" "oak" SALICACEAE ' Populus trichocarpa Torrey 6: Gray Salix sp. "willow" CRUCIFERAE Alliaria petiolata (Bieberstein) Barbarea vulgaris R. Brown Brassica sp. Descurainia sophia (Linnaeus) ERICACEAE Azalea sp. SAXIFRAGACEAE Hydrangea sp. Ribes sp. ROSACEAE Amelanchier canadensis (Linnaeus) inconspicuus inconspicuus joanae lewisi cavatus, punctulatus inconspicuus, joanae abnormis, bardi, ptmctulatus, jocmae punctulatus terminalis lugubris, terminalis canaliculatus bardi cavatus \ terminalis canaliculatus canaliculatus Ceanothus integerrimus Hooker 6t Arnott abnormis, inconspicuus ROSACEAE (continued) C. tomentosus Parry Ceanothus sp. Cerocarpus betuloides Nuttall Crataegus douglasi Lindley Crataegus spp. Prunus serotina Ehrhart P. subcordata Bentham P. virginiana Linnaeus Prunus spp. Pyrus spp. Spiraea spp. "American plum" "apple" "choke cherry" "hawthorn" "red haw" "white thorn" "wild cherry" LEGUIINOSAE Lupinus arboreus Sims Prosopis sp. "alfalfa" inconspicuus abnormis, cavatus, incorwpicwis, punctulatus, longilobus inconspicuus longilobus elegans, canaliculatus, impres- sus, terminalis canaliculatus inconspicuus lugubris, canaliculatus bardi, impressus lugubris, canaliculatus elegans, lugubris, canaliculatus canaliculatus lugubris, cyanipennis, oanali- culatus inconspicuus elegans lugubris canaliculatus alticolus punctulatus cavatus flexiventris, joanae, lugubris, canaliculatus LEGUIINOSAB (continued) "pear: "sweet pea" "vetch" CORNACEAE Camus altemifolia Linnaeus C. racemasa Lamarck ~ Camus spp. "dogwood" VITACEAE "grape" HIPPOCAS'I‘ANACEAE H ippacastaneum sp. "buckeye" ANACARDIACEAE Rhus diversiloba Torrey 6: Gray R. ovata S. Watson UMBELLIFERAE Conium maculatum Linnaeus Heracleum lanatum Michaux Heracleum sp. Lomatium sp. Pastinaca sativa Linnaeus SOLANACEAE Solarium sp. inconspicuus oregonus lugubris elegans, terminalis term inalis terminalis impressus cavatus lugubris, canaliculatus inconspicuus inconspicuus punctulatus vittatus, ptmctulatus inconspicuus inconspicuus elegans punctulatus CONVOLVULACEAE "morning glory" POLEMONIACEAE Palemanium reptans Linnaeus HYDROPHYLLACEAE Phacelia tanacetifalia Bentham Phacelia sp. CAPRIFOLIACEAE Viburnum lentaga Linnaeus Viburnum sp. COMPOSI'I‘AE Achillea millefalium var. lanulasa (Nuttall) Ambrosia trifida Linnaeus Artemesia sp. Baccharis pilularis de Candolle Balsamarhiza sagittata Nuttall Conyza sp. Manalapia lancealata Nuttall M. major de Candolle Senecia aureus Linnaeus S. serra Hooker Senecia sp. Silphium perfaliatum Linnaeus Taraxacum afficinale Wiggers "composite" incanspicwis terminalis rusticus rusticus, jahnsonarum lugubris, canaliculatus elegans bardi labiatus punctulatus punctulatus vittatus labiatus jahnsanarum rusticus, inconspicuus lugubris vittatus bardi labiatus canaliculatus punctulatus, longilabus GRAMINEAE Bramus inermis Leysser "glam" LILIACEAE Smilacina racemasa (Linnaeus) Smilax ecirrhata (Engelmann) S. herbacea Linnaeus S. hispida Muhlenberg Trillium grandiflarum (Michaux) Veratrum califarnicum Durand V. viride Aiton IRIDACEAE Iris sp. elegans abnormis, inconspicuus, oregonus bardi, punctulatus, langilobus lugubris lugubris lugubris lugubris lugubris picipennis, longilabus lugubris lugubris