This work is licensed under the Creative Commons Attribution-Noncommercial-Share Alike 3.0 United States License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-sa/3.0/us/ or send a letter to Creative Commons, 171 Second Street, Suite 300, San Francisco, California, 94105, USA. * • •&;:'W'&;:&yf.-} . Elaphrus olivaceus LeConte. 1863. Female, dorsal aspect. Length of body 7.6 mm. Illustration by D. R. Maddison. THE GENERA OF HOLARCTIC ELAPHRINI AND SPECIES OF ELAPHRUS FABRICIUS (COLEOPTERA: CARABIDAE): CLASSIFICATION, PHYLOGENY AND ZOOGEOGRAPHY.' Henri Goulet Biosystematics Research Institute Agriculture Canada K. W. Neatby Bldg., Ottawa, Ontario K1A 0C6 Quaestiones Entomologicae 19:219-482 1983 ABSTRACT The tribe, genera, and subgenera of Elaphrini are redefined on characters of adults and larvae. Recognized are three genera of Elaphrini, (Diacheila Motschulsky, Blethisa Bonelli, and Elaphrus Fabriciusj, four subgenera of Elaphrus Fabricius, (Arctelaphrus Semenov, Neoelaphrus Hatch, Elaphrus, and Elaphroterus Semenov), 34 species and 3 subspecies of Elaphrus. Keys to genera of Elaphrini and to subgenera, species and subspecies of Elaphrus are given for adults and known larvae. Four species are described as new: E. lindrothi (type locality: United States: Illinois, Jackson Co., 3 mi. N. Pomona), E. marginicollis (type locality: United States: Colorado, Jack's Gulch, Roosevelt N.F.), E. mimus (type locality: United States: California, Angwin), and E. comatus (type locality: China, Heilung Kiang, Harbin). The following synonymies are proposed for the first time: Elaphrotatus Semenov 1895 = Elaphroterus Semenov 1895; Elaphrus ruscarius foveatus Pierce 1948 = Elaphrus finitimus Casey 1920; Elaphrus clairvillei lynni Pierce 1948 = Elaphrus clairvillei Kirby 1837. Treatment of each species includes: synonymic list, diagnostic combination and description of adults and larvae, discussion of variation, derivation of the specific epithet, geographic distribution, collecting notes, taxonomic notes, and geographical affinities. Important character states are illustrated. Geographical distributions are mapped for all North American species of Elaphrus . Results of statistical analyses of geographic variation of each species are discussed. Relationships of genera and subgenera of Elaphrini are established using separate procedures of phenetic and cladistic systematics, based independently on characters of adults and larvae. A phytogeny is reconstructed for genera of Elaphrini, and for subgenera and species o/Elaphrus based on structural characters of adults and larvae. It is postulated that the ancestral elaphrine stock evolved and radiated in tropical Asia where it became extinct except for the immediate ancestor of the elaphrines surviving in the temperate zone of northernmost Siberia and Alaska in the Late Cretaceous. There, it radiated and gave rise to ancestors of extant genera and subgenera. The history of elaphrine evolution is a succession whereby ancestral peripheral elements extend into areas of low diversity followed by radiation. This pattern was repeated with the formation of the cold temperate, 'Modified and expanded from a thesis submitted to the University of Alberta in partial fulfillment of the requirements for the degree of Doctor of Philosophy. 220 Goulet boreal and arctic zones. RESUME A I'aide des caracteres morphologiques de I'adulte et de la larve, la tribu, genres et sous-genres des Elaphrini sont definis de nouveau. Nous reconnaissons trois genres a I'interieur de la tribu (Diacheila Motschulskay, Blethisa Bonelli et Elaphrus Fabricius), quatre sous-genres a I'interieur du genre Elaphrus (Arctelaphrus Semenov, Neoelaphrus Hatch, Elaphrus, et Elaphroterus Semenov), ainsi que 34 especes et 3 sous-especes du genre Elaphrus. Nous presentons des clefs de determination pour les adultes et larves connues des genres d'Elaphrini, ainsi que des sous-genres, especes et sous-especes cTElaphrus. Quatre nouvelles especes pour la science sont decrites: E. lindrothi (localite-type: Etats Unis: Illinois, Comte de Jackson, 5 km au nord de Pomona) E. marginicollis localite-type: Etats Unis: Colorado, Jack's Gulch, Foret Nationale de Roosevelt), E. mimus (localite-type; Etats Unis: Californie, Angwin), et E. comatus (localite-type: Chine: Heilung Kiang, Harbin). Les synonymes suivants sont proposes pour la premiere fois: Elaphrotatus Semenov 1895 = Elaphroterus Semenov 1895; Elaphrus ruscarius foveatus Pierce 1948 = Elaphrus finitimus Casey 1920; Elapharus clairvillei lynni Pierce 1948 = Elaphrus clairvillei Kirby 1837. Pour chaque espece traitee dans ce travail, les informations suivantes sont incluses: liste des synonymes, diagnose et description de I'adulte et de la larve, discussion de la variation geographique, origine des noms nouveaux proposes, repartition geographique, notes sur I'habitat et la biologic, notes taxonomiques et affinites geographiques. Les caracteres morphologiques importants sont illustres de meme que la repartition geographique des especes nearctiques du genre Elaphrus. Les resultats de Vanalyse statistique de la variation geographique de chaque espece sont egalement discutes. Les relations d'affinite entre les genres et sous-genres d'Elaphrini ont ete etablies a partir des techniques phenetiques et cladistiques, basees independamment sur les caracteres morphologiques de I'adulte et de la larve. Nous presentons egalement un arbre phylogenetique des genres d'Elaphrini et des sous-genres et especes ^"Elaphrus etabli a I'aide des caracteres de I'adulte et de la larve. Nous croyons que la lignee ancestrale des Elaphrini s'est developpee et repandue en Asie tropicale pour ensuite y disparaitre sauf pour Vancetre immediat des Elaphrini qui a probablement survecu dans les regions temperees de la Siberie septentrionale et de VAlaska a la fin du Cretace. Cet ancetre par la suite a evolue dans cette region et donne naissance aux lignees ancestrales des genres et sous-genres actuels. L'histoire evolutive des Elaphrini est percue comme une sussession d'invasions d'elements peripheriques vers des regions de faible diversity suivie de speciation. Ce patron s'est repete lors de la formation des regions temperees, boreales et arctiques. TABLE OF CONTENTS Introduction 221 Materials and Methods 222 Materials 222 Methods 225 Tribe Elaphrini 230 Key to Genera of Elaphrini 232 Genus Diacheila Motschulsky 235 Genus Blethisa Bonelli 236 Genus Elaphrus Fabricius 238 Key to Subgenera of Elaphrus Fabricius 239 Subgenus Arctelaphrus Semenov 241 Subgenus Neoelaphrus Hatch 246 Key to Species of Subgenus Neoelaphrus Hatch 247 Subgenus Elaphrus Fabricius 282 Key to Species of Subgenus Elaphrus Fabricius 284 Subgenus Elaphroterus Semenov 322 Key to Species and Subspecies of Subgenus Elaphroterus Semenov 323 Notes on Structures Correlated with Cryptic Coloration of Adults of Elaphrus with Their Substrate 375 Genera of Holarctic Elaphrini and Species of Elaphrus 221 Phenetics and Cladistics: Larvae and Adults 376 Phylogeny of Elaphrini 445 Classification of Elaphrini 451 Zoogeography of Elaphrini 452 Concluding Remarks 465 Acknowledgements 465 Literature Cited 466 INTRODUCTION Since capturing my first specimen of Elaphrus in 1962, I have remained excited by these beetles. Adults of most species are beautifully sculptured, and some are brilliantly coloured. Moreover, the marked specialization in habitat requirements of many species fascinated me (Goulet, 1964). Adults of Elaphrus are easy to recognize because of their cicindeloid shape, and four rows of large elytral depressions (pits). Unfortunately many species of the subgenus Elaphrus are difficult to characterize. However, Lindroth, (1961) in his revision of North American species, laid the groundwork for further studies. This work is intended as a continuation of Lindroth's work. I deal with intraspecific variation, larvae, behaviour, and habitat requirements. Although I focus much of my efforts on North American species, I include all Palaearctic taxa known to me. I gathered large amounts of structural evidence about adults and larvae to test separate phylogenetic reconstructions for congruence, and to help students of fossil insects working with fragments of specimens. More detailed descriptions are in my thesis (1978, University of Alberta, Edmonton, Canada). Finally, I attempt to trace past zoogeographical events. Cicindela riparia Linnaeus , 1758, was the first formally recognized species of Elaphrus. Fabricius (1775) erected the genus Elaphrus to include the above species, and others that are today in Notiophilus Dumeril, 1806, and Bembidion Latreille, 1802. Latreille (1810) designated E. riparius as type species, and excluded Bembidion from Elaphrus. Dejean (1826) published the first revision and restricted Elaphrus to its present concept. Some authors after Dejean used the genus Elaphrus in a wider sense: Brulle(1834) included Pelophila Dejean, 1828, and Blethisa Bonelli, 1810; Lacordaire (1854) added Opisthius Kirby, 1837. However, Dejean's concept became generally accepted. Semonov (1895, 1926), who was studying the rich Russian Elaphrus fauna, recognized the natural species-groups of Elaphrus, and arranged the species in five subgenera. Larvae were first described by Schiodte (1867). Major advances in knowledge of larvae were made by van Emden (1919, 1942), Lindroth (1954) and Luff (1976). Presently all elaphrine genera and subgenera can be recognized in larval stages. In a few recent works, precise habitats of many species were described