ZooKeys 245: 1–1722, doi: 10.3897/zookeys.245.3416
Catalogue of Geadephaga (Coleoptera, Adephaga) of America, north of Mexico
Yves Bousquet 1
1 Agriculture and Agri-Food Canada, Central Experimental Farm, Ottawa, Ontario, Canada

Corresponding author: Yves Bousquet (yves.bousquet@agr.gc.ca)

Academic editor: Terry Erwin

received 24 May 2012 | accepted 19 September 2012 | Published 28 November 2012

(C) 2012 Yves Bousquet. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

For reference, use of the paginated PDF or printed version of this article is recommended.


All scientific names of Trachypachidae, Rhysodidae, and Carabidae (including cicindelines) recorded from America north of Mexico are catalogued. Available species-group names are listed in their original combinations with the author(s), year of publication, page citation, type locality, location of the name-bearing type, and etymology for many patronymic names. In addition, the reference in which a given species-group name is first synonymized is recorded for invalid taxa. Genus-group names are listed with the author(s), year of publication, page citation, type species with way of fixation, and etymology for most. The reference in which a given genus-group name is first synonymized is recorded for many invalid taxa. Family-group names are listed with the author(s), year of publication, page citation, and type genus. The geographical distribution of all species-group taxa is briefly summarized and their state and province records are indicated.

One new genus-group taxon, Randallius new subgenus (type species: Chlaenius purpuricollis Randall, 1838), one new replacement name, Pterostichus amadeus new name for Pterostichus vexatus Bousquet, 1985, and three changes in precedence, Ellipsoptera rubicunda (Harris, 1911) for Ellipsoptera marutha (Dow, 1911), Badister micans LeConte, 1844 for Badister ocularis Casey, 1920, and Agonum deplanatum Ménétriés, 1843 for Agonum fallianum (Leng, 1919), are proposed. Five new genus-group synonymies and 65 new species-group synonymies, one new species-group status, and 12 new combinations (see Appendix 5) are established.

The work also includes a discussion of the notable private North American carabid collections, a synopsis of all extant world geadephagan tribes and subfamilies, a brief faunistic assessment of the fauna, a list of valid species-group taxa, a list of North American fossil Geadephaga (Appendix 1), a list of North American Geadephaga larvae described or illustrated (Appendix 2), a list of Geadephaga species described from specimens mislabeled as from North America (Appendix 3), a list of unavailable Geadephaga names listed from North America (Appendix 4), a list of nomenclatural acts included in this catalogue (Appendix 5), a complete bibliography with indication of the dates of publication in addition to the year, and indices of personal names, supraspecific names, and species-group names.


Ground beetles, Trachypachidae, Rhysodidae, Carabidae, North America


The Adephaga, a name coined by the Swiss entomologist and botanist Joseph Philippe de Clairville [1742-1830] in 1806, represents the second largest suborder of Coleoptera with an estimated 39, 300 species described to 2005. The group is undisputedly natural, based on the presence of several synapomorphies in the adult and immature stages (Beutel and Ribera 2005: 53; Beutel et al. 2008; Lawrence et al. 2011). The term Adephaga comes from the Greek word adephagos meaning gluttonous, greedy, in reference to the predaceous habits of adults and larvae of the vast majority of the species. Conventionally the Adephaga are divided into two groups, the Geadephaga for the terrestrial families and the Hydradephaga for the aquatic families.

The extant hydradephagan families include the Gyrinidae (about 875 species), Haliplidae (about 220 species), Noteridae (about 250 species), Amphizoidae (five species), Hygrobiidae (six species), Dytiscidae (about 3, 700 species), Aspidytidae (two species), and Meruidae (one species). Some studies, based on structural features of the adult (Burmeister 1976; Baehr 1979) and larva (Ruhnau 1986) as well as molecular data (Shull et al. 2001; Ribera et al. 2002; Hunt et al. 2007), suggest that the Hydradephaga is monophyletic. Other studies, including recent DNA sequence analyses (Maddison et al. 2009), indicate a polyphyletic origin for the complex.

The extant geadephagan groups include the trachypachids (six species), rhysodids (about 355 species), cicindelids (about 2, 415 species), and carabids (about 31, 490 species). The monophyletic origin of the Geadephaga was supported in some structural and molecular studies but rejected in others (see Maddison et al. 2009 for an overview). While the taxonomic concept of the hydradephagan families is stable, that of the geadephagan families is not. Several authors consider either the trachypachids, rhysodids, or cicindelids as Carabidae.

This work catalogues all geadephagan taxa of America, north of Mexico. The last catalogue covering the Geadephaga of the region is that of Bousquet and Larochelle in 1993. Since then relatively few taxonomic studies have been published on the North American fauna. The increased interest toward the inadequately known but amazingly rich Neotropical Region is probably one of the reasons behind the situation. So, is there a need for this catalogue? For one, it is more informative than the previous one. It includes, besides the usual information on nomenclature, the type locality of each available species, locations of the primary type specimens, references to the original synonymies of invalid names, and a short description of the geographical distribution of each species. Furthermore, a number of errors were discovered in the previous catalogue and needed to be corrected.

Brief history

The first checklist / catalogue covering the North American Geadephaga was the checklist of beetles of the United States by Friedrich Ernst Melsheimer [1784-1873] published in July 1853. The interest for this work originated with the establishment in 1842 of the first entomological society in America, The Entomological Society of Pennsylvania. The compilation of this list was one of the main objects of the Society (Sorensen 1995: 17) and it prevailed upon Melsheimer, the first and only President of the Society, to complete the task. The manuscript was delivered in 1848 to the Smithsonian Institution in Washington. Its secretary, Joseph Henry, asked Samuel S. Haldeman and John L. LeConte to advise on its publication. The two gentlemen volunteered to update the manuscript, which delayed its release considerably. The work was a straightforward list of valid species, with abridged references and synonyms but without distributional data, arranged under the valid generic names. Although limited to the United States, it included more than 90% of the species known from North America at the time. Melsheimer, a physician by profession, was the son of Frederick Valentine Melsheimer [1749-1814] who in 1806 published the first book on American entomology, a 60-page booklet entitled “Catalogue of insects of Pennsylvania. Part first.” It enumerates 111 genera and 1, 363 species of Coleoptera (Meisel 1929: 367), though almost all of them are nomina nuda.

In April 1866, John Lawrence LeConte [1825-1883] published the first part of a checklist of the Coleoptera of North America (north of Mexico) for the Smithsonian Institution. It covered the Adephaga and a large section of the Polyphaga. The first 49 pages, which included the Adephaga, were reprinted with minor modifications from a list already issued in March 1863. The list included synonyms but no geographical information. The second part of the checklist, covering the Chrysomeloidea and Curculionoidea, was never published. Two additional checklists of North American beetles would be published in the United States during the xix Century, both straightforward lists without geographical data. The first one, issued in 1874, was authored by George Robert Crotch [1842-1874], a British coleopterist who at the time was assistant to Hermann Hagen at the Museum of Comparative Zoology. A supplement to Crotch’s checklist was authored in 1880 by Edward Payson Austin, an amateur coleopterist and member of the Cambridge Entomological Club in its early years. The second checklist was published in 1885 by Samuel Henshaw [1852-1941], then assistant to Professor Hyatt at Lowell Technological Institute. Three supplements, in 1887, 1889, and 1895, were later issued by Henshaw.

In Europe, the German Max Gemminger [1820-1887] and Freiherr Edgar von Harold [1830-1886] published, between 1868 and 1876, a checklist of beetles of the world in 12 volumes, compiling 77, 008 species over 3, 800 pages. The Geadephaga were included in the first (Carabidae including cicindelids and trachypachids), second (paussids on pages 700-706), and third volumes (rhysodids on pages 867-868), all issued in 1868. Along with each specific name the authors listed the publication year as well as the original reference and region(s) of capture. This work spurred a large number of additions and corrections by many coleopterists. It stood alone in its class until the publication of the Coleopterorum Catalogus under the editorship of Walther Junk and Sigmund Schenkling. Published between 1909 and 1940, this catalogue was issued in 170 parts forming 30 volumes and involved the participation of more than 60 entomologists. A list by parts and another by families can be found in Blackwelder (1957: 1022-1034). The Geadephaga were covered in parts 1 (Rhysodidae by Raffaello Gestro in 1910), 5 (Paussinae by R. Gestro in 1910), 86 (Cicindelinae by Walther Horn in 1926), 91, 92, 97, 98, 104, 112, 115, 121, 124, 126, and 127 (Carabidae, including trachypachids, by Ernst Csiki between 1927 and 1933). Second editions of the Rhysodidae, by Walter D. Hincks in 1950, and Paussinae, by Emile Janssens in 1953, were issued much later.

While the Coleopterorum Catalogus was being published in Berlin, Charles William Leng [1859-1941], then director of the museum at the Staten Island Institute of Arts and Sciences, released in 1920 his catalogue of the Coleoptera of America, north of Mexico, still known as the “Leng catalogue.” His goal was “to enumerate systematically all the species of Coleoptera described prior to January 1, 1919 ... with consecutive numbers, synonyms, citation of original description, and an indication of distribution.” Leng and Andrew J. Mutchler in 1927 (covering the years 1919-1924) and 1933 (for 1925-1932), Richard E. Blackwelder in 1939 (for 1933-1938), and Blackwelder and his wife, Ruth M. Blackwelder, in 1948 (for 1939-1947) published supplements to Leng’s catalogue.

In 1972, Ross H. Arnett, Jr. [1919-1999], the catalyst behind the birth of the Coleopterist’s Society and its journal The Coleopterists Bulletin, initiated the “North American beetle fauna project” (NABF) with the help of a small group of coleopterists. The main goal of this cooperative adventure was to “produce a series of manuals for the identification of the species of beetles of the United States and adjacent Canada, and adjacent Mexico.” Although no such book was ever published, a preliminary checklist of North American beetles, known as the “Red Version, ” was compiled by 1976 by Richard E. Blackwelder and Arnett. This version was used as a “working copy” for the next one, the “Yellow Version” defined as the “definitive checklist and the one which will be kept up-to-date.” Of this version, only two families would be compiled and published (July 1977), the Cupedidae by Arnett and the Carabidae (including trachypachids but excluding cicindelids) by Terry L. Erwin, Donald R. Whitehead, and George E. Ball. The “Red Version” was reissued with modifications in 1983 under the editorship of Arnett.

In November 1978, the Science and Educational Administration, USDA, released its first fascicle, covering the family Heteroceridae, of “A catalogue of the Coleoptera of America north of Mexico.” The goal was to “supplant the Leng catalogue and supply additional essential information.” A total of 34 fascicles, treating various family-group taxa, would be published up to February 1997. Among the fascicles, one only, the Rhysodidae by Ross T. Bell in 1985, deals with Geadephaga.

In 1993, Bousquet and Larochelle published the first catalogue specifically devoted to the geadephagan beetles of North America. They listed, for the first time, the original combination of every available species-group taxon and provided a general idea of the distribution of each species by listing state and province records. One of the goals behind their work was to stimulate interest toward publication of distributional records as done regularly in Europe.

In 1998, Wolfgang Lorenz issued the first edition of his “Systematic list of extant ground beetles of the world” compiling 32, 567 species (in 1861 genera) of Geadephaga. Despite being limited to scientific names with their authors and publication years, the list soon became a useful tool to those interested in carabids. A second edition was released in 2005, compiling the same information for 34, 281 extant species, placed in 1929 genera.

The first catalogue of the world Coleoptera published is that of Schönherr issued in four parts, 1806, 1808, 1817 and 1826. The Carabidae were grouped in the following genera: Scarites (23 species), Cychrus (seven species), Manticora (two species), Carabus (340 species), Calosoma (12 species), Galerita (nine species), Brachinus (16 species), Anthia (27 species), Agra (three species), Collyris (four species), Odocantha [sic!] (seven species), Drypta (four species), Cicindela (67 species), Elaphrus (11 species), Scolytes [sic!] (three species), all included in the first volume (1806), and Paussus (ten species) and Cerapterus (two species) included in the third volume (1817). Overall 547 species of Geadephaga were listed along with references and synonyms. By comparison, the number of Carabidae (including Cicindelinae) listed in the four catalogue editions of the Dejean collection amounted to 104 (first edition, 1802), 908 (second edition, 1821), 2494 (third edition, 1833), and 2791 (fourth edition, 1836).

A comparison of the number of valid species and genera between this and previous checklists / catalogues is presented in Table 1.

Table 1.

North American Geadephaga species/genera counts in checklists.

Publications Trachyp Rhysod Cicindel Carabid Total
Melsheimer 1853 0 3/1 64/4 935/112 1002/117
LeConte 1866 2/1 2/2 65/4 1090/107 1159/117
Gemminger & Harold 1868 2/1 2/1 61/5 1167/124 1232/131
Crotch 1874a 2/1 2/2 67/4 1097/118 1168/125
Henshaw 1885 2/1 4/2 70/4 1179/114 1255/121
Leng 1920 2/1 4/2 114/4 2207/207 2327/214
Coleopterorum Catalogus 1926-33 6/1 4/2 70/4 2916/144 2996/151
Erwin et al. 1977 3/1 9/21 109/42 2308/169 2429/176
Bousquet & Larochelle 1993 3/1 8/2 107/4 2230/183 2348/190
Present catalogue 3/1 8/2 112/12 2316/193 2439/208

1 Species count from Bell (1985b)

2 Species count from Boyd (1982)

Nomenclatural and distributional information

The information on species-group taxa comprises a nomenclatural and a distributional component. The nomenclatural component consists of the scientific name with its author, date and page of publication, the type locality (see section Type locality under “Nomenclature” below), and the repository of the name-bearing type of each valid and invalid taxon. In addition, the reference in which a given scientific name is first synonymized is listed. Such references were difficult to find for several names, simply because they were never compiled before. Taxa listed as varieties subsequently to their original descriptions were not considered as listed in synonymy but those listed as aberrations or as “simple varieties” were. Codens used for collection repositories are given in the next section. When available, the accession numbers of name-bearing types for each institution are recorded.

This catalogue deals with extant available taxa. Fossil taxa are listed in Appendix 1. Unavailable names found in the literature are listed in Appendix 4 without comment. Listings of valid species-group names are alphabetic but listings of invalid names are chronologic. Synonyms of adventive and Holarctic species found in North America are selective. Misidentifications by subsequent authors are not listed. All species-group names are given in their original combinations.

The distributional component consists of a list of state and province records, using the same two-letter postal service style abbreviations used in the 1993 catalogue (Table 2), and a short description of the distribution, usually referring to the northeasternmost, northwesternmost, southwesternmost, and southeasternmost states or provinces. In addition, records for Cape Breton Island, the Queen Charlotte Islands, Vancouver Island, and the Channel Islands are indicated in parentheses after their respective provinces or states. Western Hemisphere countries are listed for species found south of the area covered. States and provinces placed in quotation marks in the descriptive section indicate that only the state or province was given without further precision in the reference cited. The starting point for the distributional records used in this work is Bousquet and Larochelle’s (1993) catalogue. However, many of their records were undocumented or came from old lists and were not always reliable. State and province records undocumented or considered doubtful are shown in square brackets following the accepted records. Except for the Amara records which come from identifications generally made by Fritz Hieke, almost all records from CMNH specimens are based on identifications made by Robert L. Davidson, those from LSAM specimens on identifications made by Igor Sokolov, and those from CNC, MCZ, and USNM specimens from identifications or confirmations made by myself. The records provided by Ken Karns and Brian Raber are based on identifications made by Robert L. Davidson.

Table 2.

Two-letter abbreviations for political regions covered by this catalogue.

AB Alberta MA Massachusetts OH Ohio
AK Alaska MB Manitoba OK Oklahoma
AL Alabama MD Maryland ON Ontario
AR Arkansas ME Maine OR Oregon
AZ Arizona MI Michigan PA Pennsylvania
BC British Columbia MN Minnesota PE Prince Edward Island
CA California MO Missouri PM St.Pierre and Miquelon
CO Colorado MS Mississippi QC Quebec
CT Connecticut MT Montana RI Rhode Island
DC District of Columbia NB New Brunswick SC South Carolina
DE Delaware NC North Carolina SD South Dakota
FL Florida ND North Dakota SK Saskatchewan
GA Georgia NE Nebraska TN Tennessee
GL Greenland NF Newfoundland TX Texas
IA Iowa NH New Hampshire UT Utah
ID Idaho NJ New Jersey VA Virginia
IL Illinois NM New Mexico VT Vermont
IN Indiana NS Nova Scotia WA Washington
KS Kansas NT Northwest Territories WI Wisconsin
KY Kentucky NU Nunavut WV West Virginia
LA Louisiana NV Nevada WY Wyoming
LB Labrador NY New York YT Yukon Territory

The information on supraspecific taxa consists of the scientific name with its author and date and page of publication. Type species of genus-group taxa are also given, in their original combinations, followed by the valid names in parentheses when applicable, and type genera are listed for family-group taxa. Etymology is given for all valid generic names and for some of the invalid names; the works of Brown (1956) and Cailleux and Komorn (1981) have been particularly useful.

The listing of valid supraspecific taxa is “phylogenetic, ” starting with taxa putatively branching off early along the evolutionary path of the group. Synonyms of supraspecific taxa are listed chronologically. If readily available, the first reference in which a given genus-group name is synonymized is included.

In the references section, titles of journals are cited in full. Titles of papers and books using alphabets other than Latin have been translated into English and the original language listed in square brackets after the title. An improvised title is given in square brackets, in the language used by the author(s), to papers without formal title. Unless otherwise noted, all references listed were seen. Except when only the year was found, the date of publication [DP] is given in square brackets at the end of each citation.

Institution / collection acronyms and abbreviations

Collections cited in the catalogue are referred to by the abbreviations listed below.

ALM Alabama Museum of Natural History, Tuscaloosa, Alabama, USA

AMNH American Museum of Natural History, New York, New York, USA

ANSP Academy of Natural Sciences, Philadelphia, Pennsylvania, USA

BMNH The Natural History Museum, London, United Kingdom

BYUC Brigham Young University, Provo, Utah, USA

CAS California Academy of Sciences, San Francisco, California, USA

CMC Cincinnati Museum of Natural History, Cincinnati, Ohio, USA

CMN Canadian Museum of Nature, Gatineau, Quebec, Canada

CMNH Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA

CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Ontario, Canada

CUIC Cornell University Insect Collection, Cornell University, Ithaca, New York, USA

DAPC Darren A. Pollock collection, Eastern New Mexico University, Portales, New Mexico, USA

DEI Institute für Pfanzenschutzforschung (formerly Deutsches Entomologisches Institut), Kleinmachnow, Eberswalde, Germany

EMEC Essig Museum of Entomology Collection, University of California, Berkeley, California, USA

ETHZ Entomologisches Institut, Eidgenössische Technische Hochschule, Zürich, Switzerland

FFPC Foster Forbes Purrington collection, The Ohio State University, Columbus, Ohio, USA

FMNH Field Museum of Natural History, Chicago, Illinois, USA

FSCA Florida State Collection of Arthropods, Gainesville, Florida, USA

GNM Göteborgs Naturhistoriska Museum, Göteborg, Sweden

HMUG Hunterian Museum, University of Glasgow, Glasgow, United Kingdom

INHS Illinois Natural History Survey, Champaign (Urbana), Illinois, USA

IRSN Institut Royal des Sciences Naturelles, Brussels, Belgium

IZWP Museum and Institute of Zoology of the Polish Academy of Sciences, Warszawa, Poland

KSUC Kansas State University, Manhattan, Kansas, USA

LACM Los Angeles County Museum of Natural History, Los Angeles, California, USA

LMMC Lyman Entomological Museum, McGill University, Macdonald Campus, Sainte-Anne-de-Bellevue, Quebec, Canada

LSAM Louisiana State Arthropod Museum, Baton Rouge, Louisiana, USA

LSL Linnean Society, London, United Kingdom

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA

MHNG Muséum d’Histoire Naturelle, Geneva, Switzerland

MHNP Muséum National d’Histoire Naturelle, Paris, France

MSB Museum of Southwestern Biology, University of New Mexico, Albuquerque, New Mexico, USA

MSNG Museo Civico di Storia Naturale, Genoa, Italy

MSNM Museo Civico di Storia Naturale, Milano, Italy

MSNT Museo Civico di Storia Naturale, Trieste, Italy

MSUE Michigan State University, East Lansing, Michigan, USA

MVM Museum Victoria, Melbourne, Australia

NCSU North Carolina State University, Raleigh, North Carolina, USA

NHMW Naturhistorisches Museum Wien, Wien, Austria

NIAS National Institute for Agro-environmental Sciences, Tsukuba, Japan [formerly National Institute of Agricultural Sciences, Tokyo]

NMNS National Museum of Nature and Science, Tokyo, Japan

NMP National Museum, Prague, Czech Republic

NRSS Naturhistoriska Riksmuseet, Stockholm, Sweden

NSNH Nova Scotia Museum of Natural History, Halifax, Nova Scotia, Canada

ODAC Oregon Department of Agriculture, Plant Division, Salem, Oregon, USA

ORUM Collection Ouellet-Robert, Université de Montréal, Montreal, Quebec, Canada

OSAC Oregon State Arthropod Collection, Oregon State University, Corvallis, Oregon, USA

OSUO Ohio State University, Columbus, Ohio, USA

PMNH Peabody Museum of Natural History, Yale University, New Haven, Connecticut, USA

PURC Purdue State University, West Lafayette, Indiana, USA

SIM Staten Island Museum, Staten Island, New York, USA

SMEK Snow Museum of Entomology, University of Kansas, Lawrence, Kansas, USA

SMTD Staatliches Museum für Tierkunde, Dresden, Germany

TAMU Texas A&M University, College Station, Texas, USA

TMB Magyar Természettudományi Múzeum, Budapest, Hungary

TME Texas Museum of Entomology, Pipe Creek, Texas, USA

UAIC University of Arkansas, Fayetteville, Arkansas, USA

UASM Strickland Museum, University of Alberta, Edmonton, Alberta, Canada

UBC Spencer Entomological Museum, University of British Columbia, Vancouver, British Columbia, Canada

UCD University of California, Davis, California, USA

UCM University of Colorado Museum, Boulder, Colorado, USA

UICU University of Illinois, Urbana, Illinois, USA

UMAA University of Michigan, Ann Arbor, Michigan, USA

UMM Philipps-Universität Marburg, Zoologische Sammlung, Marburg, Germany

UMO The University Museum, University of Oxford, Oxford, United Kingdom

UMSP University of Minnesota, Saint Paul, Minnesota, USA

USMT Ueno Science Museum, Tokyo, Japan

USNM National Museum of Natural History, Smithsonian Institute, Washington, DC, USA

USS University of Sydney, Sydney, Australia

UZIU Uppsala Universitet, Zoologiska Museum, Uppsala, Sweden

VMNH Virginia Museum of Natural History, Martinsville, Virginia, USA

WSU Washington State University, Pullman, Washington, USA

ZILR Zoological Institute, Academy of Sciences, Saint Petersburg, Russia

ZMH Zoologiska Museum, University of Helsinki, Helsinki, Finland

ZMHB Zoologisches Museum, Humboldt Universität, Berlin, Germany

ZMLS Zoological Museum, Lund University, Lund, Sweden

ZMMU Zoological Museum, Moscow University, Moscow, Russia

ZMUA Zoologisch Museum, Universiteit van Amsterdam, Amsterdam, The Netherlands

ZMUC Zoologisk Museum, Universitets Copenhagen, Copenhagen, Denmark

ZMUO Zoological Museum, University of Oslo, Oslo, Norway

ZMUT Zoological Museum, University of Turku, Turku (= Åbo), Finland

Besides those used for provinces and states (see Table 2), the following abbreviations are used in the text:

B.P. Before Present

CAN Canada

CBI Cape Breton Island

CHI Channel Islands (Santa Barbara Islands)

DEN Denmark

DP Date of publication

FRA France

ICZN International Commission on Zoological Nomenclature

QCI Queen Charlotte Islands

USA United States of America

VCI Vancouver Island

In addition, the International Commission on Zoological Nomenclature is sometimes abridged to “Commission” and United States of America to “United States.”

Geographical terms

For simplicity, North America, north of Mexico, is referred to simply as North America in the text. Middle America refers to Mexico and the republics of Central America taken collectively. The West Indies refers to the Greater and Lesser Antilles and the Bahamas. The North American continent proper is referred to as North and Middle America.

For practical reasons, the zoogeographical regions of the world are defined following national boundaries as much as possible. The Nearctic Region corresponds to Canada, the continental United States, Saint-Pierre and Miquelon, and Greenland. Although the region extends into Mexico, its southern limit is difficult to define and often varies depending on the group under study. This concept implies that North America and the Nearctic Region are equivalent in this work. The Neotropical Region comprises Middle America and South America. The Afrotropical Region consists of Africa, including Madagascar and a number of smaller islands of the Indian Ocean, such as the Comoros, the Mascarene Islands, and the Seychelles, and of the Atlantic Ocean, such as Cape Verde Islands and São Tomé, but excludes the northern countries of Morocco (including Western Sahara), Algeria, Tunisia, Libya, and Egypt west of the Suez Canal, and the Canary and Madeira Islands. The limits of the Palaearctic Region are similar to those used in the Catalogue of Palaearctic Coleoptera (Löbl and Smetana 2003: 8). The region thus comprises Europe, Africa north of the Sahara, and Asia as far south as the Arabian Peninsula, Pakistan, Jammu and Kashmir, Himachal Pradesh, Uttar Pradesh, Nepal, Sikkim, Bhutan, Arunachal Pradesh, China, and Taiwan. The Oriental Region is Asia south of the regions used to define the southern limit of the Palaearctic Region. It includes all the Malay Archipelago (except New Guinea). The Australian Region comprises Australia, New Zealand, New Guinea, and some smaller islands of the Pacific, such as Fiji, New Britain, New Caledonia, and Solomon Islands.

The New World consists of the Nearctic, Neotropical, and Australian Regions combined and the Old World of the Oriental, Palaearctic, and Afrotropical Regions grouped. The Northern Hemisphere is the Nearctic and Palaearctic Regions combined and the Southern Hemisphere is the Afrotropical, Oriental, Australian, and Neotropical Regions united. The Western Hemisphere consists of the Nearctic and Neotropical Regions and the Eastern Hemisphere of the Palaearctic, Afrotropical, Oriental, and Australian Regions. Far East used in reference to the Palaearctic Region includes the Russian Far Eastern Region, the Korean Peninsula, Japan, Taiwan, and China excluding the Autonomous Regions of Inner Mongolia, Sinkian Uighur, and Tibet. Middle East is used for the southwestern Asian countries, including Egypt, Turkey, Syria, Lebanon, Israel, Jordan, Saudi Arabia, Yemen, Oman, Iraq, Iran, Afghanistan, and Pakistan.

The adjective “Holarctic” is used to denote a taxon that occurs naturally in both the Nearctic and Palaearctic Regions. The adjective “Australian” (as in “Australian species”) refers to the zoogeographical region, not to the country itself. The adjective “worldwide” is used to denote a genus-group or family-group taxon represented by at least one native species in all six zoogeographical regions as defined above including both the European and Asian parts of the Palaearctic Region. The adjective “endemic” indicates that the taxon is found only in the region listed.

Names of geographical places are given in their current English forms based on Merriam-Webster’s Geographical Dictionary, third edition (1997).


The rules outlined in the fourth edition of the International Code of Zoological Nomenclatural, published in 1999, have been followed throughout this catalogue. The following are comments about some nomenclatural issues.

Principle of priority. Priority for identical taxa made available the same year, whether under the same name or not, is determined by the date, other than the year, of publication. If not specified in the work itself, the publication date is the earliest day or month on which the work is demonstrated to be in existence (ICZN 1999: Article 21.3). When both works are published or assumed to be published the same day, precedence is determined by the First Reviser (Article 24.2). Unless listed in the work itself, dates of publication besides the year can be demonstrated only for some works. Those without specific dates are listed as published the last day of the year (Article 21.3.2) and priority goes to the work with a “demonstrated” date of publication. However, the situation is subject to change with new bibliographic discoveries, which could challenge the validity of synonyms (as well as relative precedence of homonyms and validity of nomenclatural acts) and bring nomenclatural instability. In this catalogue, priority was given to the publication “in prevailing usage” when the dates of publication were determined from external sources.

New taxa. In the xviii and first half of the xix Century it was common practice for authors not to indicate the attribution of the new species-group taxa. Instead, some authors added the word mihi after the specific name, usually to indicate a taxon that the author, himself, was describing. Several collectors provided names for their specimens, even for undescribed ones, and these specimens often circulated among European coleopterists through exchange, gift, or sale. Many undescribed species were subsequently described or illustrated under the collector’s names by different authors. For these, citations are provided in this catalogue only to the first description or illustration of each species unless the term “new species” or an equivalent expression (such as an asterisk preceding the specific epithet as in Say 1823a1) was included with the species-group name subsequently described or illustrated. Sometimes a species was described / illustrated by different authors the same year under the same names. One example concerned several species (i.e., Patrobus foveocollis, Patrobus fossifrons, Pterostichus adstrictus, Pterostichus ventricosus, and Pterostichus pinguedineus)2 described by Eschscholtz in 1823 in the Mémoires de la Société Impériale des Naturalistes de Moscou (volume 6) and illustrated by Fischer von Waldheim on plates available the same year (Sherborn 1922: liii), but included in his Entomographie de la Russie (volume 2) issued in 1824. In such cases, citations are given for the oldest description / illustration (for exceptions see previous entry, Principle of priority) but references to subsequent descriptions / illustrations are noted after the entry of the valid name.

New taxa first published as synonyms. The International Commission on Zoological Nomenclature admits the availability of taxa first published in an available work as junior synonyms and adopted before 1961 as valid taxa or treated as senior homonyms (ICZN 1999: Article 11.6.1). In such cases the taxa date from their first publication as synonyms. Even though this ruling has existed since the publication of the ICZN first edition in 1960, it has rarely been enforced in the carabid literature. A few cases were found during the preparation of this catalogue. For example, Notiophilus sylvaticus has been credited in the past to Eschscholtz (1833: 24) but the name was first proposed as a junior synonym of Notiophilus biguttatus Fabricius by Dejean (1831: 589). The name is credited to Dejean (1831) in this catalogue. It is possible that other cases like this one will eventually be found.

Lectotype. Prior to 2000, a lectotype could be selected by using the term “the type” instead of “lectotype” (ICZN 1999: Article 74.5). The words “type” and “holotype” are also acceptable if the author unambiguously selects a particular syntype to act as the unique name-bearing type of the taxon. This is the case for almost all designations using the word “type” or “holotype” relating to North American Carabidae published after 1950, in particular by George E. Ball and his students. In this catalogue the expression “lectotype [as type]” or “lectotype [as holotype]” applies to such cases. Unfortunately the Commission does not mandate the addition of “lectotype” labels to selected specimens, which often creates ambiguity when authors fail to do so.

Type locality. According to the ICZN (1999: Article 76.1), the type locality is the geographical place of capture of the primary type (holotype or lectotype). In the absence of a primary type, the type locality encompasses the localities of all the syntypes (Article 73.2.3). This information can be obtained from labels attached to primary types or to syntypes or from the original publication (referred to as “original citation” in the text) whichever is more inclusive, or inferred from the title of the publication or even from the name of the species. When a neotype is designated, its place of capture becomes the type locality (Article 76.3) even if the specimen was collected outside the original area. In this catalogue, type localities taken from labels or from original publications are listed as indicated although the order of the elements is sometimes changed; any additional information is placed in square brackets. Many species described in the xviii and xix Centuries had but little informative place of origin, such as a country, state, province, or large geographical area (e.g., Rocky Mountains or Lake Superior). Lindroth (1961-1969) restricted the type locality of several of these North American species by selecting a specific locality or a county within the original region specified. This practice is followed in this catalogue and specific type localities are selected for several species-group taxa. Of course, only localities where a given species was actually collected can be selected.

Notable private carabid collections

Many North American species of carabids described in the xix and beginning of the xx Centuries were from specimens held in private collections. The whereabouts of these collections are important to taxonomists. Some of the more significant ones are discussed.

Pierre François Marie Auguste Dejean (1780-1845) Collection

Dejean, a French military officer by profession, certainly held the largest private beetle collection of his time, which he built through exchanges, purchases, gifts, and his own collecting in various parts of Europe. He described a total of 289 new carabid species-group taxa from North America, of which 182 (63%) had not been described earlier according to the present catalogue. At the sale of his collection in 1840, the carabid section (which also included the agyrtid genus Pteroloma) was the most significant, not only because it contained 3, 014 species and 17, 914 specimens, but because it was the only one to include name-bearing types. Dejean did not describe a new species-group taxon during his lifetime that he did not consider a carabid. Dejean’s carabid collection (including tiger beetles) was purchased for 7, 000 francs by Marquis F. Thibault de LaFerté-Sénectère who sold it, along with his own carabids, to Baron Maximilien de Chaudoir [q.v.] in 1859. Dejean’s carabid specimens are at MHNP today. Lindroth (1955b) discussed the name-bearing types and status of almost all North American species described by Dejean.

Thomas Say (1787-1834) Collection

Say was the first naturalist born in North America to describe new species of beetles from this continent. In the course of 17 years (1817-1834), he described 164 carabid species from North American material which he believed were new to science. Based on their current status, 142 (87%) had effectively not been previously described. Say left his collection by verbal bequest through his wife to the Academy of Natural Sciences in Philadelphia in 1834 (Weiss 1936: 277). After his death, which occurred in October of the same year, the collection was shipped from New Harmony, Indiana, to Philadelphia through New Orleans. In 1836, Charles Pickering sent Say’s insects to Thaddeus W. Harris in Cambridge, Massachusetts, in order to “put them in good order, and return them in a condition to be preserved” (Harris to D.H. Storer, 2 November 1836). In the same letter Harris reported “They [Say’s specimens] arrived about the middle of July; but on examination were found to be in a deplorable condition, most of the pins having become loose, the labels detached, and the insects themselves without heads, antennae and legs, or devoured by destructive larvae, and ground to powder by the perilous shakings which they had received in their transportation from New Harmony.” In a letter to C.J. Ward, dated 8 March 1837, Harris wrote “I assure you that Mr. Say’s cabinet does not contain one half of the species which he has described; of the insects in it, many are without names, and all more or less mutilated, and so badly preserved that most of them are now absolutely worthless.” On July 16, 1838, Harris indicated in a letter to S.G. Morton (see Fox 1902: 11) that he had “been obliged to bake a considerable part of the insects lately belonging to Mr. Say twice, and some of them three times, in order to destroy the vermin with which they are infested.” Say’s collection was returned to the Academy of Natural Sciences in Philadelphia in March 1842 “in such a state of ruin and dilapidation as to be almost useless” (Ruschenberger 1852: 25).

During his life, Say sent some of his specimens abroad including many to Dejean in Paris (see Dejean 1826: vi). Fortunately Dejean’s carabid collection has remained intact and in good condition to this day. In their attempt to bring taxonomic stability to Say’s names, Lindroth and Freitag (1969) selected lectotypes for eight carabid species described by Say for which Say’s authentic specimens could be located in Dejean’s collection. They also designated neotypes from the MCZ material for 131 of the remaining 156 of Say’s species leaving the tiger beetles (14 species) and a few taxa, all currently considered invalid, without type specimens. Say’s species were interpreted by Lindroth and Freitag from LeConte’s concept according to his collection. LeConte never saw Say’s collection and his interpretation of Say’s species came exclusively from the original descriptions which he considered adequate: “The entire destruction of his [Say’s] original specimens would be the subject of much greater regret, were it not for the fact that his descriptions are so clear as to leave scarcely a doubt regarding the object designated. I am thus enabled to assign to nearly all of his Coleoptera their proper place in the modern system” (LeConte 1859d: vi).

Thaddeus William Harris (1795-1856) Collection

Harris, well known for his work in economic entomology (his profile having appeared on every cover of the Journal of Economic Entomology for more than 35 years), described 28 new carabid species from North America. Ten (36%) are considered valid in this catalogue. To his defence, several of his species were made available by the posthumous publication of some of his letters several decades after they were written. At Say’s suggestion, Harris sent his entire collection to Thomas Say in Philadelphia, in 1825, who labeled the specimens as well as he could. Harris’ collection, which included “4, 838 specimens in 2, 241 species of Coleoptera, ” contained “many typical specimens described by Harris, Say, and others” (Scudder 1860: 72). It was bought by friends in 1858 and presented to the Boston Society of Natural History. Harris’ collection was transferred to the Museum of Comparative Zoology at Cambridge in April 1941 (Darlington 1941b: 273) where it stands separately from the general collection in two standard 25 drawer cabinets.

Gustav Graf von Mannerheim (1797-1854) Collection

Mannerheim, a Finnish noble by birth and wealthy by inheritance, described 72 new North American carabid species, all from Alaska and California. Of these, 23 (32%) had not been described previously. Mannerheim never visited the New World and his descriptions were based on specimens brought back chiefly by Russian collectors such as Johann F. Eschscholtz, Eduard L. Blaschke, Egor L. Tschernikh, and Il’ia G. Vosnesensky. His library and personal collection, which consisted, at the end, of 18, 000 species and nearly 100, 000 specimens, were sold for the sum of 8, 000 silver rubles by his widow, Countess Eva Mannerheim, in 1855 to the University of Helsinki. The money used to buy the collection came from a loan made by the Emperor to the University with the understanding that the University will pay back annually the sum of 500 rubles to the Imperial Bank of Finland which will use it for poor- and workhouses in the country (Rein 1857). Mannerheim’s collection is kept separately at the University of Helsinki (Silfverberg 1995: 43).

Jules Antoine Adolphe Henri Putzeys (1809-1882) Collection

Putzeys described 38 new North American species of carabids; 15 (39%) are listed as valid in this catalogue. He worked in close collaboration with Chaudoir, the leading carabidologist of the time, and described several new species from specimens in Chaudoir’s collection. These specimens are now in MHNP. He also gave many of his own types to Chaudoir. His personal collection was bequeathed in 1885 to the Société Royale Belge d’Entomologie under the care of the Musée Royal d’Histoire Naturelle in Brussels. Putzeys’ collection consisted of 26, 429 specimens of carabids (including cicindelids) and 6, 123 species (Preudhomme de Borre 1885: clx) as well as many other beetles and various insects.

Victor de Motschulsky (1810-1871) Collection

Motschulsky, a Russian Imperial Army Colonel, described 121 new geadephagan species from North America; 27 (22%) were undescribed at the time based on current practice. A large part of this material came from a 10-month trip he made in 1853-54 to the United States and Panama. He collected at several locations including New York, Niagara Falls, Cleveland, Cincinnati, Cawington, Lexington, the Mammoth Cave, Nashville, Louisville, New Orleans, Mobile, Montgomery, Atlanta, Washington, D.C., and Philadelphia. In the last city, he visited LeConte, Haldeman, Melsheimer, and Zeigler. The first three gentlemen gave Motschulsky several specimens from their collections including “types” (Motschulsky 1856: 16). LeConte also identified part of the beetles Motschulsky collected in Louisiana, Alabama, Georgia, and Carolina. Motschulsky’s main collection, which included almost 60, 000 specimens and about 4, 000 types of beetles, was bequeathed to the Société Impériale des Naturalistes de Moscou. It was stored in poor condition and suffered considerable damage before it was acquired in 1911 by the Zoological Museum, Moscow Lomonosov State University (Antonova 1991: 72). Keleinikova (1976) catalogued the carabid syntypes of Motschulsky’s collection at ZMMU.

Samuel Stehman Haldeman (1812-1880) Collection

Haldeman described 45 new carabid species from North America; 22 (49%) had not been described previously. In 1869 Haldeman, who had purchased Hentz’s collection, sold his collection of beetles to Simon Snyder Rathvon of Lancaster, Pennsylvania, “for about what the cases cost” (Rathvon in Geist 1881: 125). Rathvon’s collection and library were purchased for $1, 000 by Henry Bobb of East Greenville, Pennsylvania, and presented to the Franklin and Marshall College in Lancaster, Pennsylvania, as a memorial of his son (Dubbs 1903: 369). In a letter dated April 1875 and addressed to Alexander Agassiz (see below), John L. LeConte stated that he owned “all the unique types” of Haldeman. This leads one to speculate that Haldeman, a close friend of LeConte, gave his name-bearing specimens to LeConte prior to selling his collection to Rathvon.

Maximilien Stanislavovitch Baron de Chaudoir (1816-1881) Collection

Russian aristocrat of French origin, Chaudoir was not the typical insect collector. He made a single extensive collecting trip in his life, a 40 day-journey to the Caucasus in company of M.H. Hochhuth in 1845. His collection was mostly built through purchases and gifts. The single most significant purchase was LaFerté-Sénectère’s carabid collection in 1859 which included Dejean’s original specimens. In January 1874 Chaudoir gave his tiger beetle specimens, representing 713 species, to MHNP. After his death in May 1881 his collection passed into the hands of René Oberthür in Rennes as agreed upon between Chaudoir and the Oberthür brothers. Over nearly five decades, Chaudoir described 126 new carabid species based on specimens collected in North America; 58 (46%) had not been described earlier based on this catalogue.

René Oberthür died in April 1944 and his collection, certainly one of the two largest private beetle collections ever built, was classified as “monument historique” in January 1948 by the French government. The collection, which included at least five million specimens, was acquired for the sum of 32 million francs by the Muséum d’Histoire Naturelle in Paris (MHNP) in 1951 (Cambefort 2006: 249).

Henry Ulke (1821-1910) Collection

Although Ulke described only two North American carabids in his life, Bembidion nevadense in 1875 and Pterostichus johnsoni in 1889, his collection, which he sold in 1900 to the Carnegie Museum in Pittsburgh, was used extensively by LeConte and Horn and contains numerous syntypes of new species described by the two coleopterists. However, recognition of many of these syntypes can be difficult. Sometimes all syntypes were retained by LeConte and Horn while on other occasions all or some of them were returned to Ulke. Furthermore, syntypes returned to Ulke were often reincorporated in his collection with others of the same species from the same place. Usually these were marked with a number or colored square, but since many syntypes were left unmarked at the time, it is sometimes impossible to recognize them at the Carnegie Museum (Robert L. Davidson pers. comm. 2008).

John Lawrence LeConte (1825-1883) Collection

LeConte is without doubt the most outstanding North American coleopterist of the xix Century, not only because he described 514 new genus-group and about 4, 730 new species-group taxa of beetles (Henshaw 1882: 270), but because he was the first to work seriously on the classification of the North American fauna. During his scientific activity, which lasted almost 40 years, he described 724 new species-group taxa of Geadephaga from North America, 439 (61%) of which were not previously described. LeConte built his collection through his own collecting but also from gifts he received and identifications he provided to many persons from whom he usually retained all or some of the specimens. There is also little doubt that his father, Major John Eatton LeConte3, left his collection to his son. Evidence supporting this can be found in LeConte (1856a: 49) when he indicated that his second specimen of Cicindela blanda “came from the old collection of my father.” LeConte was a generous man and often offered some of his specimens to visitors (such as Motschulsky [q.v.]) or sent some to acquaintances (such as Chaudoir [q.v.] and Putzeys [q.v.]) though it seems that he retained at least one specimen of each species. Unfortunately in the xix Century the type concept for species-group taxa was not developed and LeConte sometimes gave the only syntype he had in his collection and retained one or more specimens that he acquired after the original descriptions. Therefore, syntypes of some of LeConte’s species are not in his collection. Moreover, syntypes of some of his species are difficult to find in his collection. LeConte had the habit of mixing the specimens of the new species he considered later as synonyms with those of the valid species. Since many of his specimens only bear a colored disc for label, syntypes of several of his species are not readily ascertainable.

In April 1875, LeConte wrote to his friend Alexander Agassiz, director of the Museum of Comparative Zoology in Cambridge, and expressed the wish that his collection be deposited at the museum after his death4. His collection was packed and transported by his longtime friend George Horn. It now stands separate from the general collection along with that of Horn.

LeConte used small colored paper disks to indicate the provenance of his specimens. The color system used is as follows:

Pale blue Lake Superior, Canada

Pink Middle states, i.e., Maryland, Delaware, New York, New Jersey, Pennsylvania, and possibly also Connecticut and Rhode Island

Pale pink Vermont, New Hampshire, Massachusetts

White Northern and eastern states, Canada, and possibly also Alaska

Orange (brick red) Southern and Gulf states, i.e., Virginia, North Carolina, South Carolina, Georgia, Florida, Alabama, Mississippi, Louisiana, and possibly also eastern Tennessee and Arkansas

Dark red Texas

Yellow Ohio, Illinois, Indiana, Missouri, western Tennessee, Kentucky, and possibly Iowa and the southern edge of the Great Lakes

Pale green Nebraska, Kansas, North Dakota, South Dakota, Oklahoma, Colorado, Wyoming, Montana

Dark green New Mexico

Black Utah

Silver Arizona and Valley of Gila (so including also southwestern New Mexico)

Silver with edge cut Baja California, Mexico

Gold California

Dark blue Oregon, Washington

Brown Russian America, i.e., probably the region around Colony Ross, a farming community about 75 miles north of San Francisco along the coast in California, and Alaska

George Henry Horn (1840-1897) Collection

A physician by profession, Horn authored or coauthored more than 250 papers, in which he described 154 new genera and more than 1, 600 new species of beetles, including 103 North American Geadephaga. Based on the current classification, 75 (73%) of his new geadephagan species had not been described previously. His collection and library were bequeathed to the American Entomological Society, which deposited them at the Academy of Natural Sciences in Philadelphia. In October 1974, the Horn and William G. Dietz collections were delivered to the Museum of Comparative Zoology in return for the Scudder and Morse orthopteroid insects of the MCZ (Philip D. Perkins pers. comm. 2004; see Lawrence 1973: 151). Horn’s collection is preserved along with that of LeConte apart from the general collection.

Thomas Lincoln Casey (1857-1925) Collection

From 1884 to the end of his life, Casey described 1, 864 new species-group taxa of North American Geadephaga; only 307 (16%) had not been described previously based on current concepts. Still many of his remaining “valid species” have not been subsequently studied, particularly those belonging to small species of the tribe Harpalini, and a substantial proportion will certainly end up in synonymy. Furthermore, several of Casey’s species are valid simply by chance as he did not recognize or study the proper characters (such as the male genitalia) that distinguished them from their closely related taxa known at the time. His collection, consisting of almost 117, 000 specimens, including name-bearing types for more than 9, 200 species-group taxa (Buchanan 1935: 7; Blackwelder 1950: 65), was built through Casey’s own collecting and by purchases. It was bequeathed to the United States National Museum in Washington, D.C. Casey (1918: 291) stated that “about a dozen” of his types “disappeared from ... [his] collection while temporarily at the Cambridge Museum.” The syntypes of some of these species (e.g., Bembidion militare, Tachys occultator, Amara pallida, Amara ferruginea, and Amara marylandica among Carabidae) are at the MCZ. Casey did not designate holotypes as such and therefore, unless he expressly indicated in the original description that he had but a single specimen or that a lectotype had been designated, all type specimens in his collection are syntypes.

Willis Stanley Blatchley (1859-1940) Collection

Blatchley described 12 new North American carabid species; only two (17%) are considered valid in this work. His library and large insect collection, which included 470 name-bearing specimens, were given to Purdue University. Blatchley did not select type specimens in his publications but subsequently designated lectotypes [as types] for all the new species he had described (Blatchley 1930: 33-50).

Charles Frederic August Schaeffer (1860-1934) Collection

Schaeffer, curator of the insect collection at the Museum of the Brooklyn Institute of Arts and Sciences, described 30 new carabid species; 22 (73%) are still valid today. In 1929, the Brooklyn Museum transferred 37, 100 insect specimens, including many of Schaeffer’s carabid types, to the USNM (Debbie Feher pers. comm. 2008). Currently the type material of 25 (possibly 26) of Schaeffer’s species-group taxa are in the USNM. It is clear in his 1910 paper that Schaeffer was selecting one of the specimens from his series as “the type.” However he may not have labeled them as such because lectotypes have been designated for several of his new species by various authors.

Henry Clinton Fall (1862-1939) Collection

A teacher by profession, Fall owned one of the largest private collections of North American beetles toward the end of his life, with an estimated 250, 000 specimens (including those of Charles Liebeck which came to Fall in the 1930s) representing between 14, 000 and 15, 000 species or about 90% of the fauna of the time (Darlington 1940a: 46) if one excludes the “species” described by Casey. Over a period of about 40 years, Fall described 47 new North American carabid species-group taxa; 31 (66%) are still considered valid today. He left his collection, together with his correspondence, notebooks, and reprints, to the Museum of Comparative Zoology at Harvard University where his specimens are kept separately at the end of each genus in the general collection. In one of his 1910 papers, Fall designated holotypes (as “the type”) for the first time. From this publication, “type” specimens labeled as such in his collection are considered holotypes. All original specimens of his new species described prior to 1910 should be considered syntypes. Type labels on some of these specimens were probably added after the publication of the original descriptions.

Roland Hayward (1865-1906) Collection

Hayward, a member of the Boston Stock Exchange and of the Boston Society of Natural History, described 42 new species of carabids from North America, all in the tribe Bembidiini and the genus Amara. Currently 32 (76%) are considered valid. His collection, which he built through purchases, gifts, exchanges, and his own collecting in New England as well as in Colorado, Manitoba, and New Brunswick, was bequeathed to the Museum of Comparative Zoology in Cambridge. Hayward did not designate type specimens for his new species.

Edwin Cooper Van Dyke (1869-1952) Collection

Professor Van Dyke described 73 new carabid and one new trachypachid species from North America; 54 (73%) of which had not been described previously based on their current status. His collection, consisting of about 200, 000 specimens (Essig 1953: 88), was presented to the California Academy of Sciences in 1924 where the holotypes of all but three of his 74 new species of Geadephaga are currently stored.

Howard Notman (1881-1966) Collection

Notman described 38 new carabid species from North America between 1919 and 1929; 21 (55%) had not been described previously based on their current status. In 1948 he donated his entire collection to the Staten Island Institute of Arts and Sciences, where it is still today (Smetana and Herman 2001: 118). Based on Hennessey’s (1990) type catalogue of that institution, type specimens of all new species Notman collected himself, most from the Adirondacks where he owned a summer home, are in his collection in SIM (18 in total). He also described several new species from material owned by institutions, such as the USNM. Notman did not designate type specimens in his papers of 1919 and 1920 but did so after.

Classification of Geadephaga

Unfortunately, there is no consensus among coleopterists concerning the classification of Geadephaga even at the family level. Some authors rank the cicindelids, rhysodids, and trachypachids as Carabidae while others consider one, two, or all three groups as distinct families. Even the paussines are sometimes raised to family level by modern authors. At this time, I prefer to classify the Geadephaga into three families, i.e., Trachypachidae, Rhysodidae, and Carabidae.

Following Jeannel’s (1941b-1942) classification of the carabids of France, a number of authors, mostly French and Spanish taxonomists, still recognized several families of “ground beetles.” Such an approach does not add anything to the understanding of carabid evolution. It simply adds another level to the Linnaean classification. If Jeannel’s approach is followed, it could and should have an impact on the classification of the other adephagan groups, particularly the dytiscids. Since I have been under the influence of Lindroth’s work on the carabids of Canada and Alaska, Jeannel’s approach seems to me unjustified.

Following is a discussion of the family-group taxa of Geadephaga.

Family Trachypachidae. Monophyly of this family is well supported by larval and adult apomorphies (Arndt and Beutel 1995; Beutel 1994; Beutel 1998). The systematic position of this group, however, is contentious. Bell (1966b, 1967), Bils (1976), Evans (1977a, 1985), Hammond (1979), Ward (1979), Burmeister (1980), Roughley (1981), Nichols (1985c), Beutel and Belkaceme (1986), Ruhnau (1986), Beutel and Roughley (1988), Acorn and Ball (1991), Arndt (1993), Deuve (1993), Arndt and Beutel (1995), Arndt (1998), and Beutel (1998) provided or discussed elements suggesting that trachypachids are more closely related to hydradephagans or part of Hydradephaga (i.e., Dytiscoidea) than to carabids. While most authors have regarded the Hydradephaga and Carabidae as distinct phyletic lineages, Bils (1976) and Nichols (1985c) argued that the hydradephagan-trachypachid lineage may have arisen within the Carabidae. Kavanaugh (1986) reevaluated the evidence supporting relationships of Trachypachidae with Hydradephaga. He concluded that trachypachids could be the sister-group of carabids and ranked the group as a subfamily within the Carabidae. Ponomarenko (1977) also postulated, from fossil evidence, that trachypachids and carabids are sister-groups that evolved from a common eodromeid ancestor. Beutel and Haas (1996), Kavanaugh (1998: 337), Fedorenko (2009), Dressler and Beutel (2010), and Martínez-Navarro et al. (2011) found support for monophyly of a clade including trachypachids and carabids. Recent molecular studies also suggested that trachypachids are more closely related to Geadephaga than to Hydradephaga (Shull et al. 2001; Maddison et al. 2009). In addition, pygidial gland compounds in trachypachids are more similar to those known from Carabidae than from Hydradephaga (Attygalle et al. 2004: 586). In this catalogue, trachypachids are included in the Geadephaga and given family rank.

The Trachypachidae includes two extant genera: Systolosoma Solier with two species in Chile and Argentina and Trachypachus Motschulsky with four species, one in Eurasia and three in western North America.

Many putative trachypachid fossils were found in Mesozoic deposits of Asia. Ponomarenko (1977), who studied the material, included all seven genera of trachypachid fossils in a distinct subfamily, Eodromeinae. Beutel (1998: 83) pointed out that the affinities between trachypachids and eodromeines are unclear because there are no apparent synapomorphic character states between the two groups.

Family Rhysodidae. Traditionally ranked as a distinct family, rhysodids (also known as wrinkled bark beetles) have been included within the family Carabidae in recent years by several authors following evidence or discussion provided by Bell and Bell (1962), Bell (1970), Forsyth (1972), Reichardt (1977), Baehr (1979), Beutel (1990, 1992c), Yahiro (1996), Bell (1998), Liebherr and Will (1998), and others. Some authors have treated the group as a tribe related to Scaritini or Clivinini. Reichardt (1977: 393) stated that rhysodids were “closest” to salcediines and Bell (1998: 268) even suggested that the genus Solenogenys Westwood, traditionally included within the Salcediini, is the sister-group to rhysodids. Erwin (1991a: 10) on the other hand included rhysodids within his subfamily Psydrinae along with gehringiines, psydrines, moriomorphines, patrobines, trechines, zolines, pogonines, and bembidiines. Molecular data published by Maddison et al. (1999: 125) suggest that rhysodids could be the sister-group to cicindelids and that both could be closely related to the subfamily Harpalinae. Others taxonomists, however, have continued to treat the rhysodids as a distinct family. Regenfuss (1975) and Nagel (1979) suggested that the Rhysodidae could be the sister-group of the remaining Geadephaga; Deuve (1993: 100) the sister-group to the other Adephaga (with the possible exception of Gehringiinae); Beutel and Roughley (1988) the sister-group of the remaining Adephaga excluding Gyrinidae; Beutel (1992a, 1993, 1998) the sister-group to Carabidae (without trachypachids). Recently Makarov (2008) found no evidence from the larval morphology suggesting that rhysodids are specialized Carabidae. Instead rhysodid larvae share several features with those of the suborder Archostemata. At this time, I prefer to rank rhysodids as a distinct family based on tradition but also on the fact that there is no solid morphological or molecular evidence presented to date pointing out that the Carabidae (with or without trachypachids) are paraphyletic in regard to rhysodids.

About 355 species of rhysodids are currently known and are placed into seven family-group taxa, namely Leoglymmiini, Medisorini, Rhysodini, Dhysorini, Sloanoglymmiini, Omoglymmiini, and Clinidiini. These taxa are usually ranked as subtribes when rhysodids are included in the carabids. I have followed Bousquet and Larochelle (1993) in listing them as tribes. Only the last two-mentioned tribes are represented in North America.

Tribe Leoglymmiini. This tribe contains a single species, Leoglymmius lignarius (Olliff), from Australia. Contrary to other rhysodids, the minor setae on antennomeres 5-10 are arranged in broad bands encircling the distal third of the segment and the mentum is separated from the ventral lobe of the gena by a distinct suture in its anterior half.

Tribe Medisorini. A single species, Medisores abditus Bell and Bell, belongs to this tribe. The few known specimens have been found in Cape Province in the Republic of South Africa.

Tribe Rhysodini. This tribe is confined to the Eastern Hemisphere and includes about 25 species in three genera: Rhysodes Germar (two Palaearctic species), Kupeus Bell and Bell (one New Zealand species), and Kaveinga Bell and Bell (23 Australian species).

Tribe Dhysorini. This tribe includes ten species placed in three genera, Dhysores Grouvelle in Africa, Tangarona Bell and Bell in New Zealand, and Neodhysores Bell and Bell in South America.

Tribe Sloanoglymmiini. This tribe has been proposed for one species, Sloanoglymmius planatus (Lea), endemic to southeastern Australia. The genus is taxonomically isolated and its relationship to other rhysodid genera is obscure.

Tribe Omoglymmiini. This tribe includes 180 species placed in eight genera. The group is represented in all zoogeographical regions but less so in Australia, Africa, and South America (Bell and Bell 1978: 66). The two North American species belong to the subgenus Boreoglymmius Bell and Bell, of the genus Omoglymmius Ganglbauer, along with one Japanese species. According to Bell and Bell (1983: 141), the two North American species are probably more closely related to each other than either is to the Japanese species.

Tribe Clinidiini. This tribe contains about 135 species placed in the genera Clinidium Kirby, Rhyzodiastes Fairmaire, and Grouvellina Bell and Bell. The species are found in all zoogeographical regions, including Madagascar, but are absent from the African continent. The North American fauna has only six species, five in the east and one in the west, included in the subgenus Arctoclinidium Bell of the genus Clinidium. This subgenus also contains three Palaearctic species, one in Japan and two in Europe. According to Bell and Bell (1985: 77), the North American species and the Japanese one form a clade and the European species another clade. These authors also placed the Japanese species, Clinidium veneficum Lewis, as the sister-group to Clinidium valentinei Bell of eastern North America.

Family Carabidae. Monophyly of the Carabidae, as defined here, is not evident. The layout of the prehypopharyngeal setae in the larvae (Beutel 1993) and the development of antennal pubescence in the adults (Beutel 1995) have been suggested as synapomorphies for the family. However, Arndt et al. (2005: 138) considered these character states not very convincing given the variation involved in the structures. Recent molecular sequence analyses conducted by Maddison et al. (2009) found little support for monophyly of the group no matter if the trachypachids, rhysodids, and/or cicindelids were included or excluded unless the Carabidae was considered equivalent to the Geadephaga. Therefore, the Carabidae, as defined here, could be paraphyletic in regard to rhysodids, trachypachids, and possibly even to Hydradephaga.

Carabids are found on all continents, except Antarctica, and on most islands. They range from well above the arctic circle to Tierra del Fuego and South Georgia in the Southern Hemisphere. Based on Lorenz’s (2005) checklist, 33, 920 valid species are recognized.

The current classification of the Carabidae is based mainly on morphological data of adults although molecular sequence data have been used recently to discuss various aspects of carabid phylogeny. Despite several attempts there is no consensus on the classification of several subfamilies or tribes. This is particularly evident among ‘basal grade’ carabids.5

Fossils belonging to the family Carabidae are known from the early Jurassic (Ponomarenko 1977) which suggests that the family emergence dates back to the beginning of the Jurassic or the end of the Triassic (Kryzhanovskij 1983). Ponomarenko (1977) proposed two family-group taxa of Carabidae among Mesozoic fossils, the subfamily Protorabinae for five genera and the tribe Conjunctiini for two genera.

The world classification of family-group taxa, which has been adopted for the North American fauna in this catalogue, is outlined in Table 3.

Table 3.

Classification of world family-group taxa of Carabidae. Taxa represented in North America are followed by a dot.

Subfamily Nebriinae
  Tribe Pelophilini •
  Tribe Opisthiini •
  Tribe Nebriini •
  Tribe Notiokasiini
  Tribe Notiophilini •
Subfamily Cicindinae
  Tribe Cicindini
Subfamily Carabinae
  Tribe Cychrini •
  Tribe Pamborini
  Tribe Ceroglossini
  Tribe Carabini •
Subfamily Cicindelinae
  Tribe Amblycheilini •
  Tribe Manticorini
  Tribe Megacephalini •
  Tribe Cicindelini •
  Tribe Ctenostomatini
  Tribe Collyridini
Subfamily Loricerinae
  Tribe Loricerini •
Subfamily Elaphrinae
  Tribe Elaphrini •
Subfamily Omophroninae
  Tribe Omophronini •
Subfamily Migadopinae
  Tribe Amarotypini
  Tribe Migadopini
Subfamily Hiletinae
  Tribe Hiletini
Subfamily Scaritinae
  Tribe Pasimachini •
  Tribe Carenini
  Tribe Scaritini •
  Tribe Clivinini •
  Tribe Salcediini
  Tribe Dyschiriini •
  Tribe Promecognathini •
  Tribe Dalyatini
Subfamily Broscinae
  Tribe Broscini •
Subfamily Apotominae
  Tribe Apotomini
Subfamily Siagoninae
  Tribe Enceladini
  Tribe Siagonini
  Tribe Lupercini
Subfamily Melaeninae
  Tribe Melaenini
Subfamily Gehringiinae
  Tribe Gehringiini •
Subfamily Trechinae
  Tribe Trechini •
  Tribe Zolini
  Tribe Bembidiini •
  Tribe Pogonini •
Subfamily Patrobinae
  Tribe Lissopogonini
  Tribe Patrobini •
Subfamily Psydrinae
  Tribe Psydrini •
Subfamily Moriomorphinae
  Tribe Moriomorphini
  Tribe Amblytelini
Subfamily Nototylinae
  Tribe Nototylini
Subfamily Paussinae
  Tribe Metriini •
  Tribe Mystopomini
  Tribe Ozaenini •
  Tribe Protopaussini
  Tribe Paussini
Subfamily Brachininae
  Tribe Crepidogastrini
  Tribe Brachinini •
Subfamily Harpalinae
   Supertribe Pterostichitae
  Tribe Morionini •
  Tribe Cnemalobini
  Tribe Microcheilini
  Tribe Chaetodactylini
  Tribe Cratocerini
  Tribe Abacetini •
  Tribe Pterostichini •
  Tribe Zabrini •
  Tribe Metiini
  Tribe Drimostomatini
  Tribe Chaetogenyini
  Tribe Dercylini
  Tribe Melanchitonini
  Tribe Oodini •
  Tribe Peleciini
  Tribe Brachygnathini
  Tribe Bascanini
  Tribe Panagaeini •
  Tribe Chlaeniini •
  Tribe Cuneipectini
  Tribe Orthogoniini
  Tribe Idiomorphini
  Tribe Glyptini
  Tribe Amorphomerini
   Supertribe Harpalitae
  Tribe Licinini •
  Tribe Harpalini •
  Tribe Geobaenini
  Tribe Omphreini
  Tribe Sphodrini •
  Tribe Platynini •
  Tribe Perigonini •
  Tribe Ginemini
  Tribe Enoicini
  Tribe Atranini •
  Tribe Catapieseini
  Tribe Lachnophorini •
  Tribe Pentagonicini •
  Tribe Odacanthini •
  Tribe Calophaenini
  Tribe Ctenodactylini •
  Tribe Hexagoniini
  Tribe Cyclosomini •
  Tribe Somoplatini
  Tribe Masoreini
  Tribe Corsyrini
  Tribe Sarothrocrepidini
  Tribe Graphipterini
  Tribe Lebiini •
  Tribe Dryptini
  Tribe Galeritini •
  Tribe Zuphiini •
  Tribe Physocrotaphini
  Tribe Anthiini
  Tribe Helluonini •
  Tribe Xenaroswellianini
  Tribe Pseudomorphini •

Subfamily Nebriinae. This subfamily includes the tribes Nebriini, Notiokasiini, Notiophilini, Opisthiini, and Pelophilini. All but notiokasiines are Northern Hemisphere elements and represented in North America. Evidence supporting monophyly of Nebriinae is not overwhelming. The only known synapomorphy in the adult stage is the asetose parameres (Kavanaugh and Nègre 1983), a character state found in other, clearly unrelated carabid lineages. Arndt (1993: 21) listed three putative synapomorphies upon examination of the larval morphology. The molecular data analyses by Maddison et al. (1999: 125) provided only moderate support for monophyly of the subfamily and Kavanaugh’s (1998) phylogenetic analysis suggested that this subfamily represents a grade rather than a clade.

The subfamilies Nebriinae and Carabinae could be closely related as pointed out by Jeannel (1940: 7), Bell (1967: 105), Beutel (1992c: 57), and Su et al. (2004: 49). Both groups have open procoxal cavities, contrary to the remaining carabids. In addition, the external lamella of the metepimeron is completely covered and functionally replaced by an extension of the hind margin of the anepisternum (Beutel 1992c: 57). Some authors (e.g., Lorenz 2005: 125) also include the cicindines within the subfamily suggesting a close relationship between these groups. Based on similarities in the genitalia, Deuve (1993: 125) raised the possibility that the Hydradephaga, trachypachids, omophronines, and nebriines form a clade.

Tribe Pelophilini. This tribe includes a single genus, Pelophila Dejean, which has been retained in the tribe Nebriini until recently. Two species are known, both living in the boreal and subarctic regions: one is circumpolar, the other restricted to Canada and Alaska. Kavanaugh (1996: 34) suggested that the genus represents the sister-group to the remaining Nebriinae. One of Kavanaugh’s (1998: Fig. 2) cladograms suggested that Pelophila is more closely related to the tribe Nebriini than are the Opisthiini, Notiophilini, and Notiokasiini.

Tribe Opisthiini. This tribe includes two genera with five species and is doubtless monophyletic. Kavanaugh and Nègre (1983: 564) argued that opisthiines could be the sister-group to the remaining Nebriinae. On the other hand, Kavanaugh’s (1996: Fig. 1A) most parsimonious tree suggested that this tribe is the sister-group to Notiophilini and that these two tribes, along with Notiokasiini, form a clade which represents the sister-group to Nebriini.

Tribe Nebriini. This tribe contains about 600 species in the Palaearctic, Nearctic, and northern parts of the Oriental Regions. However, the group is clearly more diverse in the Palaearctic. The main genera of the tribe are Leistus Frölich, Archastes Jedlička, and particularly Nebria Latreille with more than 60% of the species. The limits of the genus Nebria are not quite settled. Kavanaugh (1995, 1996) regarded Nippononebria Uéno (including Vancouveria Kavanaugh) as the sister-group to Leistus while Ledoux and Roux (2005) listed Nippononebria and Vancouveria as subgenera of Nebria and suggested they form the sister-group to Eonebria Semenov and Znojko and Sadonebria Ledoux and Roux, a complex of 60 Palaearctic species.

Tribe Notiokasiini. This tribe contains a single species, Notiokasis chaudoiri Kavanaugh and Nègre, found in South America. Although the relationships of the tribe are obscure (Kavanaugh and Nègre 1983), Kavanaugh (1996: 33) found 12 synapomorphies supporting monophyly of a clade including notiokasiines, notiophilines, and opisthiines.

Tribe Notiophilini. The tribe includes a single genus, Notiophilus Duméril, very characteristic in the adult stage. The larvae, however, are similar in most structural features to those of Nebriini as pointed out by van Emden (1942). Jeannel (1941b: 175) included Notiophilini, Nebriini (with Pelophila), and Opisthiini in his family Nebriidae, suggesting implicitly a close relationship between the three groups. Kavanaugh’s (1996: Fig. 1A) most parsimonious cladogram suggested a sister-group relationship between Notiophilini and Opisthiini based on adult and larval morphological data. Based on confluent procoxal cavities, Nichols (1985c: Fig. 5) considered the tribe to be the sister-group to {Omophronini + Trachypachini + Hydradephaga}. Erwin (1991a: 11) noted that notiophilines, along with omophronines, hiletines, and trachypachids, have the first mesotarsomere slightly dilated and with squamate setae underneath. However, it remains to ascertain whether this character state is synapomorphic or convergent. Based on female reproductive tracts, Liebherr and Will (1998: 146) suggested that the tribe Notiophilini represents the sister-group to {Opisthiini + Nebriini (with Pelophila) + Omophronini}.

Notiophilines, with about 55 species described to date, live in the Nearctic and Palaearctic Regions and at higher altitudes in the northern parts of the Neotropical and Oriental Regions. They are more speciose in Asia than anywhere else. The phylogenetic relationships of the species have not been studied yet.

Subfamily Cicindinae. This subfamily includes two species, Archaeocindis johnbeckeri (Bänninger) from the Persian Gulf (Kuwait and Iran) and Cicindis horni Bruch from the Córdoba Province of Argentina. Very little can be said at this time about the relationships of the subfamily except that it represents a basal grade carabid taxon. Kryzhanovskij (1976a: 87) associated cicindines with paussines (excluding metriines) and nototylines; Nagel (1979, 1987) and Roig-Juñent et al. (2011) viewed them as the sister-group to paussines. Ball (1979: 100), however, doubted such proposed affinities between cicindines and paussines. Erwin (1985, 1991a), followed by Lorenz (2005: 125), included the Cicindini in the Nebriitae. Kavanaugh and Erwin (1991) studied the structural features and reviewed the relationships of the group. They concluded that cicindines are best placed in a distinct supertribe near the Nebriitae and Elaphritae (sensu Kryzhanovskij 1976a: 88). Kavanaugh’s (1998: Fig. 3) phylogenetic analysis using 153 characters of adult external and male genitalic structures suggested that cicindines may be closely related to omophronines, carabines, cychrines, and cicindelines. Aspects of the behaviour and life history of the Argentine species have been published recently (Erwin and Aschero 2004).

Subfamily Carabinae. This subfamily contains about 1, 300 species (Lorenz 2005: [i]) placed in four tribes: Cychrini, Pamborini, Ceroglossini, and Carabini. Most authors agree that the subfamily is monophyletic. According to Deuve (2004: 32), adults of this group are characterized by two significant autopomorphies: the presence of two strip-like apodemes flanking the basal orifice of the median lobe of the aedeagus and the presence of an alveolus on the epipleurite of the abdominal segment IX at the opening of the defensive gland. Arndt (1998: 179) noted several autopomorphies in larvae of Carabinae: an extensive decrease of number of setae on the tergites and sternites with an increase in the number of pores, size reduction of the sensorial appendage on antennomere III, and a markedly sclerotized body.

Tribe Cychrini. This well-defined and likely monophyletic group of about 200 species is restricted to the Northern Hemisphere. Osawa et al. (2004: 31) and Su et al. (2004: 49), based on molecular data, argued that the tribe is probably the sister-group to the remaining clades of the subfamily. Moore (1966b), Prüser and Mossakowski (1998: 316), and Arndt (1998: 180), based on morphological data, suggested that pamborines are the closest relatives to cychrines. Jeannel (1941b: 167) indicated that cychrines are more closely related to pamborines and ceroglossines than to carabines (sensu stricto) and calosomatines based on the shape of the parameres.

Relationships among the four genera have not been investigated. Whether or not Sphaeroderus Dejean and Scaphinotus Dejean, the two endemic North American genera, are sister-groups, as suggested by Erwin (2007a: 139), remains to be investigated. For example, Prüser and Mossakowski (1998: 316) listed several putative synapomorphies suggesting that Cychrus Fabricius and Sphaeroderus are sister-groups (Cychropsis Boileau was not included in their analysis). The phylogenetic tree by Osawa et al. (2004: Fig. 5.2) based on molecular sequence data suggested that Scaphinotus is the sister-group to the remaining Cychrini and Sphaeroderus the sister-group to Cychropsis.

Tribe Pamborini. The 13 species currently included in this tribe are placed in two genera: Pamborus Latreille from Australia and the monospecific Maoripamborus Brookes from New Zealand. Jeannel (1941b: 94) stated that pamborines are more closely related to ceroglossines than to any other Carabinae.

Tribe Ceroglossini. This tribe comprises only the genus Ceroglossus Solier (eight species and 46 subspecies) which is restricted to Chile and western Argentina. The genus has traditionally been included within the Carabini but recent analyses based on molecular sequence data suggest that ceroglossines are more closely related to pamborines than to carabines (Prüser and Mossakowski 1998: 297; Su et al. 2004: 50) or form the sister-group to the remaining members of Carabinae (Osawa et al. 2004: Fig. 5.2). Arndt (1998: 179) found evidence from the larval morphology to support the latter hypothesis.

Tribe Carabini. Carabines rank among the most popular groups for beetle collectors. Adults of many species are elegant, colorful, and large (often exceeding 15 mm). Such interest has generated a market for these beetles, particularly in Europe, and unfortunately also a race to describe new varieties, morphs, and aberrations. More than 1, 080 species of Carabini are recognized today worldwide. They inhabit all zoogeographical regions but are much more diverse in the Palaearctic Region than anywhere else.

The supraspecific classification of Carabini is debated. Some authors recognize only two genera, Carabus Linnaeus and Calosoma Weber, while others admit many, more or less clearly defined genera which are often grouped in two subtribes: Carabina and Calosomatina. I have followed the first approach and list all North American species in the genera Carabus (15 species) and Calosoma (41 species). The main difference between the two genera is the regression (or complete disappearance) of the ostial ligula of the aedeagus in members of Carabus (Deuve 2004: 33).

Based on morphological (larvae and endophallus of adults) and molecular sequence data, Deuve (2004) recognized eight major lineages within the genus Carabus: Spinulati, Digitulati, Lipastrimorphi, Archicarabomorphi, Tachypogenici, Metacarabi, Arcifera, and Neocarabi. The North American species are arrayed in nine subgenera: Carabus s.str. belongs to the Digitulati, Archicarabus Seidlitz to Archicarabomorphi, Tachypus Weber to Tachypogenici, Megodontus Solier to Neocarabi, and Diocarabus Reitter, Homoeocarabus Reitter, Aulonocarabus Reitter, Hemicarabus Géhin, and Tanaocarabus Reitter to Metacarabi.

Jeannel (1940: 68) recognized two major lineages within the calosomatines: the Calosomes lobés with a membranous ligula at the proximal opening of the male median lobe and the Calosomes ongulés with a chitinized ligula. The first lineage is represented in the Australian and Palaearctic Regions, and by six species belonging to the genus-group taxa Calosoma s.str. and Calodrepa Motschulsky in the Nearctic Region, the West Indies, and Mexico. The second lineage was divided by Jeannel (1940: 69-71) into three clades, the phyletic series of Castrida-Caminara represented in North America by a single species belonging to the genus-group taxon Castrida Motschulsky, the phyletic series of Callisthenes represented in the Nearctic Region by 23 species of the taxa Chrysostigma Kirby and Callistenia Lapouge, and the phyletic series of Callitropa represented in North America by 11 species of Blaptosoma Géhin, Carabosoma Géhin, Camegonia Lapouge, and Callitropa Motschulsky.

The systematic position of Aplothorax burchelli Waterhouse, a species endemic to the island of Saint Helena off the west coast of Africa, is controversial. Jeannel (1940) included the taxon within his Calosomes ongulés but Basilewsky (1972: 18-22) was convinced that the species is a relict of an old clade that evolved before the splitting of the Carabus and Calosoma lineages. He advocated placing the species in a distinct tribe which, in his opinion, was as justified as those of Pamborini and Ceroglossini.

Subfamily Cicindelinae. This group, referred to as the tiger beetles, has been regarded traditionally as a distinct family, but more and more coleopterists include it within the carabids. There is little doubt, based on characters of adults and larvae, that cicindelines form a monophyletic lineage. Relationships of the group, however, remain uncertain. It has been regarded as the sister-group to the remaining Carabidae by Nichols (1985c) and as the sister-group to Carabidae (minus paussines) by Regenfuss (1975). A close affinity between this subfamily and the Carabinae has been suggested by Erwin and Sims (1984: 366), Deuve (1993: 160; 2004: 32), Kavanaugh (1998: 338), and Liebherr and Will (1998: 151), although Liebherr and Will also emphasized that the Cicindelinae could instead be closely related to Promecognathini and Amarotypini. Maddison et al. (1999) indicated that most of their phylogenetic analyses of 18 rDNA place the Cicindelinae and Rhysodidae as sister-groups, near the Harpalinae. They also pointed out that the alternative placement of the cicindelines outside the Carabidae was more parsimonious than placing them among the basal-grade carabids. Deuve (2004: 32) noted two exceptional and primitive character states shared between cicindelines and carabines: presence of the abdominal tergite X in the male and presence of a phallobase on the aedeagus. He also pointed out numerous similarities between the two groups: the ectodermal genital ducts of the females are almost identical with a vagina differentiated in a bursa copulatrix, the presence of a sclerotized ligular apophysis, the presence of a filiform spermatheca and absence of an accessory gland, the presence of rod-shaped apophyses on the female abdominal epipleurites VIII allowing the formation of a telescopic egg-laying tube, the parameres of the aedeagus are glabrous and symmetrical, the endophallus often shows comparable dentiform sclerites, the digestion is extra-oral, and the ventral surface of the adult body often shows metallic coloration, an exceptional character state in the Adephaga.

This subfamily currently includes more than 2, 500 species distributed worldwide, except Tasmania, Antarctica, and remote oceanic islands, with the greatest diversity in the tropics (Pearson 1988). Tiger beetles are classified by most authors, following Horn (1926), into two major lineages, Alocosternales with a very narrow and deeply longitudinally-furrowed metepisternum in the adult and Platysternales with a wider metepisternum that has either no longitudinal furrow or a horseshoe-shaped furrow posteriorly. These groups are often listed as supertribes Collyriditae and Cicindelitae respectively. However, based on a combined analysis of molecular and morphological data, Vogler and Barraclough (1998: 254) noted that collyridites nested within the cicindelites, rendering the latter paraphyletic. Arndt (1998: 178) also noted that, based on larval character states, Cicindelitae and the tribe Megacephalini are not monophyletic. Based on the above information, the species of Cicindelidae are simply placed here in six tribes without further grouping.

Tribe Amblycheilini. This tribe includes the genera Omus Eschscholtz and Amblycheila Say represented in North America and Mexico, and the genus Pycnochila Motschulsky with one species in the Strait of Magellan. Amblycheilines have been classified in the past within the megacephalines but larval characters (Arndt and Putchkov 1997) and mitochondrial and nuclear RNA gene sequences (Vogler and Barraclough 1998: 251) suggest a basal position for amblycheilines, well removed from the true megacephalines. The tribe, however, may well be a grade rather than a clade. For example, Arndt (1998: 178) placed Omus as the sister-group to the remaining Cicindelinae based on larval characters.

Tribe Manticorini. This tribe includes 14 species, arrayed in the genera Mantica Kolbe (one rarely collected species from southern Namibia) and Manticora Fabricius (13 Afrotropical species). Contrary to other tiger beetles, males of this tribe have asymmetric mandibles and unexpanded protarsomeres (Werner 2000: 22).

Tribe Megacephalini. This tribe includes about 200 species arrayed in 11 genera (see Naviaux 2007: 15). Even without the amblycheilines, monophyly of the tribe is doubtful. For example, the genus Oxycheila Dejean, traditionally considered a member of Megacephalini, nested with rather strong support within the basal groups of Cicindelini in Vogler and Barraclough’s (1998: 254) cladistic analysis based on molecular and morphological data.

Tribe Cicindelini. This tribe is by far the most diversified clade of tiger beetles. The number of recognized genera varies to a great extent among taxonomists. In this work the 98 North American species (202 species-group taxa) are placed in nine genera. All but two (Cylindera and Cicindela) of these genera are New World endemics. Arndt (1998: 178) stated that Cicindelini forms the sister-group to {Ctenostomatini + Collyridini}.

Tribe Ctenostomatini. This tribe includes two genera: Ctenostoma Klug with about 115 Neotropical species, and Pogonostoma Klug with about 110 Madagascan species. Members of this tribe are synapomorphic in lacking the articulated hook at the extremity of the inner lobe of the maxilla (Jeannel 1946: 104).

Tribe Collyridini. This tribe contains about 335 species in Asia, of which one extends into the Australian Region. The species are arrayed in two subtribes: Tricondylina for the genera Derocrania Chaudoir (16 species) and Tricondyla Latreille (about 45 species), and Collyridina for the genera Protocollyris Mandl (16 species), Neocollyris Horn (about 250 species), and Collyris Fabricius (ten species). Naviaux (1994: 149) indicated the structural differences between the two subtribes.

Subfamily Loricerinae. This subfamily contains a single genus, Loricera Latreille, although some authors have treated Elliptosoma Wollaston, with one species from Madeira, as a distinct genus. The group is restricted to the Nearctic and Palaearctic Regions with some taxa found on mountains in the northern parts of the Neotropical and Oriental Regions. Loricerini is a basal grade taxon with obscure affinity. Jeannel (1941b: 80) associated loricerines with the Carabinae, Nebriinae, Cicindelinae, Elaphrinae, Omophroninae, and Siagoninae (including Promecognathus) in his Caraboidea Simplicia, characterized by the absence of metepimeric lobes (Jeannel 1941b: 93). Bell (1967: 105) included loricerines within his Anisochaeta Isopleuri along with elaphrines, scaritines, and cicindelines. Arndt (1993: 22) found several common derived larval features in larvae of Loricerinae and Cicindelinae to suggest a sister-group relationship between the two taxa. Maddison et al. (1999: 126) pointed out that placement of Loricera within {Migadopini + Amarotypini} received relatively strong support in their 18S rDNA analyses. These three taxa were also recovered as a monophyletic unit in analyses of the same gene by Ribera et al. (2005: 290). Vigna Taglianti and Rossi (1998: 515) indicated that loricerines could be closely related to elaphrines based on the presence of the same parasitic laboulbeniales species found on these groups. Erwin (2007a: 69) listed Elaphrini as the sister-group to Loricerini.

Recently Sciaky and Facchini (1999) described a new subgenus (Plesioloricera) for the new Chinese species Loricera balli which has eight, instead of 12 or more, striae. This species could possibly be the most basal taxon of the genus.

Klausnitzer (2003) described a new species, Loricera electrica, based on a larva found in Baltic amber. He believes the species probably belongs to the pilicornis group as defined by Ball and Erwin (1969).

Subfamily Elaphrinae. This subfamily includes a single tribe with obscure relationships. Bell (1967) listed the Elaphrini within his Isopleuri along with Loricerini, Scaritini, and Cicindelini. Following Jeannel’s (1941b: 214) intuition, both Kryzhanovskij (1976a: 88) and Erwin (1985: 469) considered the elaphrines as the sister-group to Migadopini (with and without Amarotypus respectively) and this hypothesis was supported by Deuve’s (1993: 160) study of the female genitalia and Roig-Juñent’s (1998: Figs 9-10) parsimony analysis using 33 adult and larval characters. On the other hand, Goulet (1983: 445) regarded Melaenini and the subtribe Broscina as the taxa most closely related to Elaphrini. This possibility was found most parsimonious by Maddison et al. (1999) based on their molecular sequence analysis which did not, however, include melaenine exemplars. In addition, Yahiro (1990: 42) reported a similar type of alimentary canal in elaphrines and broscines. Roig-Juñent’s (1998) parsimony analysis shows that elaphrines are not related to broscines. Parsimony analysis based on the female reproductive tract characters placed Elaphrini as the sister-group to {Opisthiini + Nebriini + Notiophilini + Omophronini} or near the Promecognathini and Amarotypini (Liebherr and Will 1998: 146).

The tribe is represented only in the Northern Hemisphere and includes three genera, all represented in the Nearctic Region.

Subfamily Omophroninae. This subfamily includes a single genus, Omophron Latreille. Some authors have suggested that Omophronini, Trachypachidae, and the Hydradephaga form a monophyletic group. Putative synapomorphies proposed for the complex include the presence, in the larvae, of an undivided cardo and a dorsal insertion of the femoro-tibial extensor (Ruhnau 1986) and, in the adults, the atrophy of intertergal muscle M24 (Bils 1976), partial housing of the procoxae by the mesosternum, and the prominent prosternal process contacting the metasternum (Nichols 1985c). Deuve (1993: 160) noted the presence of a peculiar sclerotized structure (named “sclerite helminthoïde”) in the female genitalia of omophronines, nebriines (sensu lato), trachypachids, and the Hydradephaga which raises the possibility of close phylogenetic affinities between these groups. Jeannel (1941b: 219) suggested that cicindelines were most closely related to omophronines. Bell (1967: 106) indicated that Omophronini might be aberrant Hemipleuri, a group including nebriines (sensu lato) and carabines (sensu lato). Kavanaugh (1998: 338), based on a parsimonious analysis of adult characters, suggested a close affinity between Omophroninae, Carabinae, and Cicindelinae. Liebherr and Will (1998: 156) listed several “potentially synapomorphic characters” supporting placement of omophronines with nebriines (sensu lato) and their preferred cladogram, based on 20 characters of female ovipositors and reproductive tracts, placed them as the sister-group to notiophilines. Many authors, however, have treated the Omophronini as a basal-grade taxon with unclear affinity. Based on larval character states, Landry and Bousquet (1984) found no evidence to indicate a sister-group to Omophronini. Such conclusions were also reached by Beutel (1991) from his study of the larval head and adult thoracic structures. Erwin (2007a: 63) indicated the possibility that cicindines were closely related to omophronines.

The genus Omophron includes about 70 species and is represented in all zoogeographical regions except the Australian one. There is no evidence yet known to suggest that the Nearctic or the Western Hemisphere species form a clade within the genus.

Subfamily Migadopinae. This subfamily currently consists of two tribes: Amarotypini and Migadopini. One of the main character states of the group is the presence of a long scutellar stria reaching the apical declivity of the elytra (Jeannel 1938b: 4) as in members of Pelophila. This characteristic, however, is absent in members of the genus Aquilex Moret, which have a short scutellar stria (Moret 2005: 30). Jeannel (1938b) revised the species of migadopines (as Migadopidae) and classified them into two groups: Monolobinae for the Chilean genus Monolobus Solier (two species) and Migadopinae for the remaining genera which are represented in South America and in the Australian Region. He also postulated that elaphrines were without doubt the group most closely related to migadopines in the Northern Hemisphere (Jeannel 1938b: 10). Moore et al. (1987: 65) included the migadopines from Australia within the supertribe Elaphritae.

Tribe Amarotypini. This tribe includes a single species, Amarotypus edwardsii Bates, from New Zealand. Until recently the species was placed in the tribe Migadopini but it differs by having a setiform unguitractor plate between the tarsal claws which is missing in migadopines. Erwin (1985: 469) postulated that Amarotypini could be the sister-group to {Migadopini + Elaphrini}. In Liebherr and Will’s (1998: Fig. 57) preferred cladogram, based on 20 characters of the female ovipositors and reproductive tract, amarotypines were positioned as the sister-group to promecognathines. In Roig-Juñent (2004: Fig. 1) phylogenetic analysis, based on 57 characters of the adult morphology, the genus Amarotypus Bates nested inside the remaining migadopine genera, as the sister-group to {Calathosoma + Stichonotus}.

Tribe Migadopini. This group of about 30 species in 15 genera is restricted to the temperate areas of the Neotropical and Australian Regions. Moret (2005: 30) recently proposed the subtribe Aquilicina for the genera Aquilex Moret (one species in Ecuador) and Rhytidognathus Chaudoir (one species in Uruguay). He also pointed out the close relationship between the South American genus Lissopterus Waterhouse (two species from the Tierra del Fuego Archipelago and Falkland Islands) and the New Zealand genera Loxomerus Chaudoir (five species) and Calathosoma Jeannel (one species). The phylogenetic analysis performed by Roig-Juñent (2004) do not support Moret’s conclusions although Aquilex was recovered as the sister-group to the other migadopine genera. Liebherr and Will (1998: 147) alluded to the possibility that the tribe is not monophyletic.

Subfamily Hiletinae. This subfamily includes two genera, Hiletus Schiødte (= Camaragnathus Bocandé) with six species in tropical Africa and Eucamaragnathus Jeannel with 15 species in the Afrotropical (six species), Oriental (five species), and Neotropical (four species) Regions. Jeannel (1941b: 80; 1946: 209) postulated that hiletines were closely related to scaritines (sensu lato) based mainly on the fact that these two taxa were the only disjuncti (i.e., with disjunct mesocoxae) with the metepimera lobed as in the conjuncti. Erwin and Stork (1985: 445) believed that hiletines were related to cnemalobines (as Cnemacanthini), elaphrines, migadopines, promecognathines, pseudomorphines, scaritines, and siagonines based on some tarsal character states and suggested that this complex forms the sister-group to the paussine-brachinine clade based on the presence of distinct epimera, brushy non-styliform parameres, long empodial unguitractor plates, and non-conjunct mesocoxae. They also concluded that hiletines represent the sister-group to {scaritines + cnemalobines (as Cnemacanthini) + pseudomorphines} and that this clade was characterized by having a single long guard seta at the apex of the fifth tarsomere which projects between the two tarsal claws.

Subfamily Scaritinae. This subfamily is inadequately defined and possibly polyphyletic. The species possess a mesothoracic peduncle which frees the prothorax from the elytra and allows greater mobility (Basilewsky 1973: 9). It includes about 1, 870 species worldwide which are grouped in this work into eight tribes: Pasimachini, Carenini, Scaritini, Clivinini, Dyschiriini, Salcediini, Promecognathini, and Dalyatini. Until the subfamily is better defined, it is difficult to comment on its relationships. Jeannel (1938a: 206) underlined a number of morphological features in the adults suggesting that scaritines and hiletines shared a common ancestor. Lindroth (1969b: xxiii) hypothesized that the similarities in “habitus and general organization” between scaritines (including promecognathines, clivinines, and dyschiriines) and broscines are probably an indication of close affinity.

Tribe Pasimachini. This tribe is represented by the genera Pasimachus Bonelli, with 32 species ranging collectively from southern Canada to Panama, and Mouhotia Laporte, with three species in eastern Asia. Monophyly of this group is doubtful. Bänninger (1950: 484) noted that if pasimachines and carenines are retained as distinct subtribes, Mouhotia cannot be placed in either of them and a separate subtribe would need to be established. To avoid proposing a new family-group name, Mouhotia is included here in the Pasimachini. Lorenz (2005: 132) included it with the carenines.

Relationships of the tribe have been little discussed. Sloane (1905b: 103) retained pasimachines and carenines under one family-group name implying a close relationship between the two groups. Nichols (1988a: 214) argued that Pasimachini is the sister-group to the tribe Carenini.

Tribe Carenini. This clade, which is endemic to Australia, includes about 195 species placed in ten genera. The genus Scaraphites Westwood (seven Australian species), usually listed as a member of this tribe (e.g., Lorenz 2005: 133), has been removed from it and placed in the tribe Scaritini by Moore and Lawrence (1994: 512). According to Moore and Lawrence (1994: 503), carenines represent the sister-group to the remaining Scaritini (sensu lato, i.e., pasimachines, scaritines, clivinines, dyschiriines).

Tribe Scaritini. This tribe, with about 495 species in 42 genera, is represented in all zoogeographical regions but is predominantly tropical. Four subtribes are currently recognized: Acanthoscelitina (one Afrotropical species in Acanthoscelis Dejean), Oxylobina (about 30 Oriental species in Oxylobus Chaudoir), Scapterina (about 25 Australian-Oriental-Afrotropical species in Passalidius Chaudoir, Scapterus Dejean, Parathlibops Basilewsky, Thlibops Putzeys, and Steganomma Macleay), and Scaritina (including the Madagascan storthodontines and dyscherines and the Afrotropical ochyropines [one species] and corintascarines [one species]), with the bulk of the species. Jeannel (1946: 220) assigned scapterines to his Clivinitae.

The North American fauna is represented by seven species of Scarites Fabricius.

Tribe Clivinini. This tribe is the most diversified group of the subfamily with about 60 genera. The inclusion of the clivinines within the Scaritinae has not been challenged often but parsimony analysis based on the female reproductive tract by Liebherr and Will (1998) suggests that clivinines could be more closely related to rhysodids than to scaritines. They also emphasized that the defensive secretions of the pygidial glands differ drastically between the two groups: clivinines use ketones or quinones while scaritines eject aliphatic acids.

At least three putative clades are recognized within the tribe and are usually ranked as subtribes. The forcipatorines, exclusively Neotropical, include the genera Camptidius Putzeys (one species), Camptodontus Dejean (14 species), Forcipator Maindron (four species), Kultianella Perrault (two species), Obadius Burmeister (two species), and Stratiotes Putzeys (two species). According to Perrault (1994: 686), members of this clade differ from those of remaining clivinines in having the ligula truncate and glabrous instead of prolonged and with at least one apical seta, the gula either vanishing posteriorly or very narrow instead of wide, the first antennomere asetose instead of having an apical seta (except in a few genera), the penultimate labial palpomere glabrous (except in one genus) instead of having two setae (except in some genera), and the clypeus glabrous (except in two species) instead of having a seta on each side. Another clade, the ardistomines, is restricted to the Western Hemisphere. Bousquet (2006c) restricted the group to the genera Ardistomis Putzeys, Semiardistomis Kult, and Aspidoglossa Putzeys whose members have a projection on pleurite VII. Kult (1950b) also included the genus Neoreicheia Kult. Whitehead (in Reichardt 1977: 386, 391) remarked that Oxydrepanus Putzeys was “doubtless related to Neoreicheia” and probably belonged to the ardistomine radiation along with such Old World genera as Reicheia Saulcy, Syleter Andrewes, and allies. Basilewsky (1973: 276) indicated that ardistomines are relatively closely related to dyschiriines. The third clade, the reicheiines, is represented only in the Old World and contains many genera including Reicheia Saulcy, Trilophus Andrewes, Typhloreicheia Holdhaus, Trilophidius Jeannel, and Leleuporella Basilewsky.

Iablokoff-Khnzorian (1960: 93) described a new genus, Dyschiriomimus, from Baltic amber which he viewed as an intermediate taxon between Dyschirius and Clivina. However, Fedorenko (1996: 37) believed the taxon is a typical clivinine more closely related to Trilophus and Oxydrepanus than to Clivina.

Tribe Salcediini. This tribe includes four genera placed in three subtribes following Bell (1998): Salcedia Fairmaire (nine Indo-African species) in Salcediina, Holoprizus Putzeys (one Amazonian species) and Solenogenys Westwood (three Brazilian species) in Solenogenyina, and Androzelma Dostal (one Vietnamese species) in Androzelmina. According to Bell (1998: 264), salcediines, forcipatorines, and clivinines form a well-defined clade supported by the following synapomorphies: the metepimeron is lobate and overlaps the anterior corners of abdominal sternum 2; the elytron possesses a ventral carina in form of a projected lobe which engages the dorsal angles of abdominal sternum VII; the labial pits on the mentum each have a posterior duct opening into the submental suture contrary to other scaritines in which the ducts open anteriad the suture and distant from it. Bell (1998: 265) also indicated that rhysodids shared three synapomorphies with salcediines (excluding Androzelma): a kind of coating on the exoskeleton, minute and retractile palpi, and a distinct lobe on which the eye is located. Furthermore among salcediines, members of Solenogenys share two synapomorphies with rhysodids: the medial margins of the ventral groove of the head are oblique, nearly straight, and meet near the “neck” condyle and the mandible has a dorsolateral lobe. Based on the above evidence, Bell (1998: 269) concluded that Solenogenys is the sister-group to rhysodids.

Tribe Dyschiriini. Relationships of this tribe within the Scaritinae are not documented. Fedorenko (1996: 37) suggested that dyschiriines share a common ancestor with clivinines but failed to disclose any characteristics that would support this claim. Jeannel (1946: 214) combined the ardistomines and dyschiriines in his Dyschiriitae.

Tribe Promecognathini. This small and well-defined tribe includes one genus with two species in western North America and four genera with six species in Cape Province in South Africa. Jeannel (1941b: 244; 1946: 206) postulated that promecognathines were closely related to siagonines without, however, offering any evidence. Lindroth (1961a: 125) and Kryzhanovskij (1976a: 88) associated promecognathines with scaritines (sensu lato) implying a close relationship between the two groups. Several apomorphic features, including details of the chaetotaxy, structure of the mouthparts and thorax, and marked similarity in their specialized way of attacking millipedes suggest that promecognathines could be closely related to peleciines. However, Straneo and Ball (1989) regarded the similarities between the two groups as evolutionary convergence.

McKay (1991) described a fossil from Cretaceous crater lake deposits at Orapa, Botswana, under the name Palaeoaxinidium orapensis, which he believed represents the sister-group to the Promecognathini.

Tribe Dalyatini. Molecular (Ribera et al. 2005) and morphological (Mateu and Bellés 2004) data suggest that the single, highly modified cave species of this tribe, Dalyat mirabilis Mateu from southeastern Spain, could be the sister-group to promecognathines.

Subfamily Broscinae. This subfamily includes a single tribe with about 290 species in 34 genera, arrayed in five subtribes (see Roig-Juñent 2000): Axonyina (five species), Broscina (about 75 species), Nothobroscina (about 90 species), Barypodina (about 25 species), and Creobiina (about 95 species). Broscines are represented in all major regions of the world, except the Afrotropical Region, but are more diverse in the Australian Region than anywhere else. They live almost exclusively in temperate areas, with only a few groups extending to the edges of the tropics. Several authors have suggested explicitly or implicitly that broscines are closely related to apotomines but Liebherr and Will’s (1998) morphological data on the female reproductive tract and the molecular sequence data provided by Maddison et al. (1999) did not support such an association. Erwin (1991a: 10) included broscines, apotomines, melaenines, and cymbionotines in his subfamily Broscinae.

Roig-Juñent’s (2000) parsimony analysis based on morphological characters of adults suggested that the three native North American genera (Broscosoma, Zacotus, and Miscodera) form a clade.

Subfamily Apotominae. This subfamily includes a single genus, Apotomus Illiger, with about 15 species in warm temperate and tropical regions of the Eastern Hemisphere and one species in Brazil. Kryzhanovskij (1976a: 88) and Moore et al. (1987: 122) associated apotomines with broscines and in several classifications these two groups are placed sequentially in the text. However, Roig-Juñent’s (1998: Fig. 10) parsimony analysis using 33 characters placed apotomines as the sister-group to melaenines (no Cymbionotum exemplars were included). Liebherr and Will (1998: 150) noted that apotomines do not have conjunct mesocoxae as in broscines and the other members of Jeannel’s Stylifera and that the placement of apotomines within the Stylifera should be rejected. They placed apotomines as a basal grade with clivinines and rhysodids but noted they could be closely related to scaritines and hiletines. Molecular data analyzed by Maddison et al. (1999: 128) did not provide support for a close relationship between apotomines and broscines.

Subfamily Siagoninae. This subfamily includes three genera, each arrayed in its own tribe: Enceladus Bonelli, Luperca Laporte, and Siagona Latreille. Relationships of the subfamily are obscure. Jeannel (1941b: 244; 1946: 206) associated siagonines with promecognathines. Erwin (1985: 467; 1991a: 9-10) listed siagonines with amarotypines, migadopines, elaphrines, promecognathines, hiletines, pseudomorphines, and scaritines (including cnemalobines, as Cnemacanthini) in his subfamily Scaritinae. The preferred cladogram of Liebherr and Will (1998: Fig. 57), based on the female reproductive tract, placed siagonines as the sister-group to the subfamily Carabinae. Some of the analyses on 18 rDNA performed by Maddison et al. (1999: 127) suggested that Siagona could be closely related to {Gehringia + Cymbionotum}. Based on a morphological study of larvae of Siagona and Enceladus, Grebennikov (1999: 9) did not find evidence to suggest a close relationship for siagonines.

Tribe Enceladini. Only the genus Enceladus belongs to this tribe, with one species found in the Guyana-Venezuelan area, possibly also in Amazonia (Reichardt 1977: 384).

Tribe Siagonini. This tribe contains only the genus Siagona with about 80 species in the Old World. Erwin (1978: 105) listed several apomorphic states shared by Siagona and Cymbionotum and stated that the two were undoubtedly closely related.

Tribe Lupercini. Two species of the genus Luperca are included in this tribe, one is found in tropical Africa, the other on the Indian subcontinent. Erwin (1978: 105) combined the genus with Siagona and Cymbionotum in his tribe Siagonini which he included in his subfamily Siagoninae along with the tribe Enceladini.

Subfamily Melaeninae. This subfamily includes one tribe with two genera: Melaenus Dejean with two species confined to the Oriental Region, Egypt, and the Afrotropical Region, excluding the Congo Basin and southern parts (Ball and Shpeley 2005: 37), and Cymbionotum Baudi di Selve with 20 species arrayed in two subgenera, Procoscinia Ball and Shpeley with two species in northern South America and Cymbionotum s.str. with 18 species confined to the warmer parts of the Old World. According to Ball and Shpeley (2005: 22), monophyly of this subfamily is indicated by the very long diverticulum of the spermathecal gland. Prior to these authors, the two genera had been variously classified. Several authors placed them in separate tribes though suggesting implicitly or explicitly that they were closely related (e.g., Erwin 1985: 469; Liebherr and Will 1998: 137). Others have separated the two rather widely. For example van Emden (1936a: 46) listed Melaenus in his Harpalinae piliferae from which the cymbionotines were excluded. Jeannel (1941b: 291-292) placed the genus Melaenus in his Psydritae along with psydrines, melisoderines (= moriomorphines), and meonines (= moriomorphines) and included Cymbionotum in a family-group taxon of its own which he considered closely related to siagonines (Jeannel 1946: 206).

Relationships of the subfamily are unclear. Liebherr and Will (1998: 150) suggested that Melaeninae could be closely related to Clivinini. Roig-Juñent’s (1998) parsimony analysis using 33 characters showed Melaenus to be the sister-group to apotomines; Cymbionotum exemplars were not included in his analysis.

This group has been reported in publications of the xix and early xx Centuries under the name Granigerini, because Graniger algirinus Motschulsky, the sole species included by Motschulsky in his new genus Graniger, was listed in synonymy with Coscinia semelederi Chaudoir (Chaudoir 1876d: 63). Because Coscinia Dejean was a junior homonym of Coscinia Hübner, Graniger Motschulsky became the valid name for this genus. However, Andrewes (1933: 3) showed that Motschulsky’s species was in fact identical with the ditomine Carterophonus femoralis Coquerel. Cymbionotum Baudi di Selve was the next available name for the species of Coscinia Dejean.

Subfamily Gehringiinae. This subfamily includes three genera placed in two subtribes: Gehringiina Darlington with a single western North American species, Gehringia olympica Darlington, and Helenaeina Deuve with four rarely collected species from Egypt, Turkey, Yemen, and Namibia placed in the genera Helenaea Schatzmayr and Koch and Afrogehringia Baehr, Schüle and Lorenz. The taxonomic position of the group is debated. Jeannel (1941b, 1946: 46) combined gehringiines with trachypachids and paussines (as Caraboidea Isochaeta) and both Lindroth (1969b) and Kryzhanovskij (1976a: 87) associated gehringiines with trachypachids. Bell (1967: 106) indicated that the form of the palpi and the anterior tibia suggest that gehringiines could be derived from the genus Tachys but he also raised the possibility that gehringiines could belong to his Hemipleuri, a group comprising the nebriines and carabines. In Nagel’s (1987: Fig. 2) cladistic analysis, gehringiines were positioned as the sister-group to {cicindines + paussines}. Beutel (1992c) indicated that the isochaetous protibia of gehringiines suggests that the group could be “an early offshoot of the metriine-paussine lineage.” In listing the tribe in his supertribe Psydritae, Erwin (1985: 468) suggested implicitly that gehringiines could be closely related to psydrines and patrobines and Lorenz (2005: 243) placed the tribe Gehringiini within the Psydrinae. Deuve (2005) made a detailed analysis of the morphology of gehringiine adults and concluded that several character states suggest “a very basal position in the phylogeny of adephagan Coleoptera” for gehringiines. However he also noted that the peculiar “abdominal type” found in the group is similar to that of the genus Cymbionotum. A close affiliation between Gehringia and Cymbionotum received support from the molecular analysis of Maddison et al. (1999). Arndt et al. (2005: 140) pointed out that the condition of the protibial spurs in gehringiines could not be unambiguously assigned to either of the two types found in other carabids. They noted that if the protibial spurs of gehringiines are considered to be of the isochaete type, then gehringiines could be the sister-group to Paussinae and if considered to be of the anisochaete type, they could be the sister-group to Nebriinae.

Subfamily Trechinae. Several authors agree that the tribes Trechini, Zolini, Bembidiini, and Pogonini are closely related and probably constitute a clade. Monophyly is supported by characteristic features of the adult morphology (Roig-Juñent and Cicchino 2001), larval morphology (Grebennikov and Maddison 2000: 226; 2005: 44), and 18S ribosomal sequence data (Maddison et al. 1999). In addition, males of Trechinae studied lack chiasmata in meiosis (Serrano 1981) contrary to most other Carabidae, a notable synapomorphy (Maddison and Ober 2011: 243). Jeannel (1941b: 299) also included mecyclothoracines in the subfamily (as Trechidae) but most recent authors place them within the Moriomorphinae. Deuve (1993: 156) included patrobines within the Trechinae.

As discussed under Patrobinae, this subfamily is probably the sister-group to patrobines.

Tribe Trechini. A relatively well-defined and very diverse group with more than 2, 500 species currently arranged in 170 genera or so. Although represented in all zoogeographical regions, the tribe is more diverse in the temperate zones than in the tropics. Many species are endogean or troglodytic and flightless. Casale and Laneyrie (1982: 7) classified the Trechini into six groups placed in two major complexes, one including cnidines, trechodines, and plocamotrechines characterized by the median lobe of the aedeagus being wide open dorsally, the basal orifice lying between two symmetric lobes, and one comprising the perileptines, aepines, and trechines with the median lobe partly closed dorsally, the basal orifice opening on the ventral surface of the basal bulb. However, this classification has been challenged in recent times. Uéno (1989: 12-13) presented arguments to combine cnidines with perileptines and Grebennikov and Maddison (2005: 46-47) found evidences in the larval characters that perileptines were closely related to trechodines. In this work, the Trechini are grouped into two subtribes: Trechodina (including perileptines, cnidines, and plocamotrechines) and Trechina (including aepines).

Based on larval character states, Grebennikov and Maddison (2005) suggested that Trechini is the sister-group to {Zolini + Bembidiini + Pogonini}. Arndt’s (1993: 33) analysis of larval characters suggested that trechines are closely related to tachyines.

The North American fauna comprises about 225 species grouped into nine genera, all belonging to the subtribe Trechina. Barr (1985a: 351) recognized four series among the North American genera, the Trechus series with Trechus, the Trechoblemus series with Trechoblemus, Blemus, Pseudanophthalmus, Neaphaenops, and Nelsonites, the Darlingtonea series with Darlingtonea and Ameroduvalius, and the Aphaenops series with Xenotrechus. The genus Pseudanophthalmus, whose members are cave inhabitants except for a few rare occurrences in forest floor humus and in abandoned coal mines, is closely related to Duvaliopsis Jeannel which includes six edaphic species in the Carpathian and Transylvanian Alps of eastern Europe. In fact, Barr (2004: 7) listed Duvaliopsis as a junior synonym of Pseudanophthalmus since both genera are not readily separable on purely morphological grounds. Xenotrechus, with two species in southeastern Missouri caves, is apparently the sister-group to the monospecific genus Chaetoduvalius Jeannel (Barr 2004: 10) of the Apuseni Mountains in the western Carpathians, Romania.

Tribe Zolini. The 57 species of this tribe are currently arrayed in ten genera and three subtribes: Zolina with 50 species in South America (genus Merizodus Solier) and the Australian Region (genera Oopterus Guérin-Méneville, Zolus Sharp, Synteratus Broun, Percodermus Sloane, Idacarabus Lea, Sloaneana Csiki, and Pterocyrtus Sloane), Sinozolina for the genus Sinozolus Deuve (six Chinese species), and Chalteniina for Chaltenia patagonica Roig-Juñent and Cicchino of Argentina. Jeannel (1962) recognized two lineages within Zolina based on structural features of the male genitalia.

Tribe Bembidiini. This relatively well-defined tribe is represented in all zoogeographical regions of the world. Adults possess characteristic subulate apical palpomeres (except in Horologion), a condition found otherwise only in gehringiines and a few trechines. Bembidiines are grouped into six subtribes: Bembidiina, with about 1, 350 species, is distributed worldwide but is more diverse in the temperate regions than in the tropics; Xystosomina is represented in the New World and tropical Australia (Erwin 1994: 560) by about 125 species with only one (Mioptachys flavicauda) found in North America; Tachyina (including lymnastines) with nearly 800 species is also worldwide but, contrary to Bembidiina, is more diverse in the tropics; Anillina with about 375 minute, apterous, and blind species is distributed in all zoogeographical regions; Horologionina with a single cave-inhabiting species, known only from the holotype collected in West Virginia; and Lovriciina represented by four cavernicolous species, placed in three genera (see Giachino et al. 2011), found in the Balkans. Erwin (1982b: 459) postulated that anillines and horologionines represent a grade of several lineages derived from Paratachys Casey and allies, a hypothesis refuted by Maddison and Ober (2011: 249). Arndt (1993: 33) found a number of putative synapomorphies in larvae of Tachyina and Trechini and suggested that the two taxa are sister-groups. Grebennikov (2002) and Grebennikov and Maddison (2005), working with larvae, found evidence suggesting that Anillina is the sister-group to {Tachyina + Xystosomina} and that xystosomines are probably nested within the tachyines. Van Emden (1936a) suggested that Horologion Valentine was closely related to psydrines and Jeannel (1949b: 93) believed it could be related to patrobines.

Tribe Pogonini. This tribe is found in all zoogeographical regions of the world but is more diverse, both in terms of species and lineages, in the Palaearctic Region. All 83 species currently recognized are more or less halobiont and live along sea coasts or near salt lakes. Jeannel (1941b: 552) stated that this group is related to mecyclothoracines (currently placed in the subfamily Moriomorphinae) of the Hawaiian islands and the Australian Region. Müller (1975) postulated that Pogonini is the sister-group to Bembidiini. Based on karyotypic data, Serrano and Galián (1998: 196) suggested that pogonines are closely related to Bembidiina. Arndt (1993: 33), working on larval characters, suggested a close relationship between pogonines and Bembidiini (excluding tachyines).

Subfamily Patrobinae. This subfamily, which includes the tribes Lissopogonini and Patrobini, is considered to be the sister-taxon to Trechinae by several authors based on male tarsal structure (Müller 1975), larval characteristics (Arndt 1993: 32), and similar abdominal morphology (Deuve 1993). This association is also supported by molecular sequence data (Maddison et al. 1999: 128; Maddison and Ober 2011: 243). Erwin (1985: 469) and Baehr (1998: 363) suggested that patrobines may be closely related to Moriomorphinae. Jeannel (1941b: 80-81) placed patrobines in his Limbata Stylifera along with apotomines, broscines, psydrines, moriomorphines, melaenines, trechines, bembidiines, pogonines, and zolines but indicated that some character states, particularly of the larvae, suggest that they may belong to the Limbata Conchifera.

This subfamily is found in the Northern Hemisphere and Oriental Region. Only the tribe Patrobini is represented in North America.

Tribe Lissopogonini. This tribe includes a single genus, Lissopogonus Andrewes, with eight species in Asia. The genus was originally described in the tribe Pogonini and subsequently transferred to the tribe Patrobini by Zamotajlov and Sciaky (1996: 40). Bousquet and Grebennikov (1999: 11) alluded to the possibility that Lissopogonus could be a highly derived taxon related to Patrobus and Platypatrobus based on the shared apomorphic condition of the median sulcus of the pronotum being wide and deep in the basal fifth and reaching the basal edge. Deuve and Tian (2002: 30) suggested that the genus could belong at the base of the Trechinae and Patrobinae lineages (their Trechidae).

Tribe Patrobini. The 215 species or so listed in this tribe are currently arrayed in four subtribes: Deltomerina with the genus Deltomerus Motschulsky only, Deltomerodina with the genus Deltomerodes Deuve, Patrobina with 18 genera, and Platidiolina with Platidiolus Chaudoir. In a cladistic analysis conducted by Roig-Juñent and Cicchino (2001: Fig. 1), this tribe is positioned as the sister-group to Amblytelini (currently included in the Moriomorphinae).

Subfamily Psydrinae. Following Maddison and Ober (2011: 237), this subfamily is restricted to the tribe Psydrini and includes only six species. Two (Laccocenus ambiguus Sloane and Laccocenus vicinus Moore) lives in southeastern Australia, another one (Psydrus piceus LeConte) ranges widely across the northern parts of North America, extending southwards to the mountains of northern California, Arizona, and New Mexico, and the other three, all members of the genus Nomius Laporte, are restricted to central Africa and Madagascar (two species) or to the Northern Hemisphere although apparently extinct in Asia (Nomius pygmaeus Dejean). Baehr’s (1998: Fig. 1) preliminary cladistic analysis using 19 characters of adults suggested that Psydrini could be the sister-group to {Patrobinae + the remaining Psydrinae [= Moriomorphinae]}. Relationships among the three genera of Psydrini have not been investigated.

Subfamily Moriomorphinae. Members of this subfamily were traditionally included in the Psydrinae but recent morphological (Baehr 1998) and molecular data (Maddison and Ober 2011: 237) studies suggest that the Moriomorphinae form a clade and that the group is not closely related to the true Psydrinae. Baehr (1998: 363) argued that Patrobinae could be the sister-group to Moriomorphinae. Many moriomorphines are similar to pterostichines in body form but the presence of a scrobal seta and setose parameres in almost all moriomorphines, unlike pterostichines, suggest that they are probably not closely related. Ober’s (2002) phylogenetic analysis based on molecular sequence data suggested that the subfamily Moriomorphinae, termed “austral psydrines, ” could be the sister-group to {Brachininae + Harpalinae}.

This subfamily, which includes about 470 species, is represented only in the Southern Hemisphere and is particularly diverse in the Australian Region. Five tribes were traditionally recognized (see Baehr 2004): Mecyclothoracini with about 285 species placed in the genera Neonomius Moore and Mecyclothorax Sharp; Meonini with about 20 species in the genera Raphetis Moore, Meonis Laporte, Selenochilus Chaudoir, and Meonochilus Liebherr and Marris; Moriomorphini with six species in five genera, all endemic to southeastern Australia; Tropopterini with about 50 species in seven genera; and Amblytelini with six genera and about 95 species endemic to Australia, including Tasmania. Recently, Liebherr (2011) proposed an entirely new classification, dividing the moriomorphines into two groups based on characters of the parameres. His classification is adopted here.

The genus Bembidiomorphum Champion (two species in Chile), included in this group since van Emden (1936a: 51), belongs to the Broscini (Roig-Juñent et al. 2008: 212).

Tribe Moriomorphini. This group includes about 55 species, all endemic to the Australian Region, placed in 13 genera: Celanida Laporte (one species), Melisodera Westwood (three species), Molopsida White (28 species), Moriodema Laporte (two species), Moriomorpha Laporte (one species), Neonomius Moore (three species), Pterogmus Sloane (one species), Rhaebolestes Sloane (two species), Rossjoycea Liebherr (one species), Sitaphe Moore (eight species), Teraphis Laporte (six species), Theprisa Moore (three species), and Trephisa Moore (one species). These species are characterized by having elongate, parallel-sided parameres that are glabrous or sparsely clothed with very short setae.

Tribe Amblytelini. This tribe contains about 415 species in 12 genera: Amblytelus Erichson (43 species), Dystrichothorax Blackburn (48 species), Epelyx Blackburn (five species), Mecyclothorax Sharp (about 280 species), Meonis Laporte (16 species), Meonochilus Liebherr and Marris (six species), Paratrichothorax Baehr (one species), Pseudamblytelus Baehr (one species), Raphetis Moore (three species), Selenochilus Chaudoir (six species), Trichamblytelus Baehr (one species), and Tropopterus Solier (four species). These species are restricted to Australia and New Zealand except those of Mecyclothorax which occur also in New Guinea, Borneo, Java, and the Polynesian islands in the Pacific Ocean and Tropopterus which are found in Chile and Peru. Amblytelines differ from members of Moriomorphini by having more setose parameres that are either shorter, basally broader and narrowly rounded apically or elongate with whiplike apex.

Subfamily Nototylinae. This subfamily includes a single species, Nototylus fryi (Schaum), known only from the female holotype collected in the state of Espírito Santo, Brazil, in the xix Century. The species is aberrant structurally: it lacks the grooming structures of the protibiae present in all other Geadephaga except Paussini and lacks the pubescence on antennomeres 5-10 which is present in other Geadephaga except Trachypachidae, Rhysodidae, and Gehringiinae (Deuve 1994b: 141). Bänninger (1927) suggested that Nototylus Gemminger and Harold was related to Ozaenini, Kryzhanovskij (1976a: 87) associated it with paussines (excluding metriines) and cicindines, and Erwin (1979: 591) postulated that the species was an independently adapted myrmecophile from an ozaenine stock. However, Ball (1979: 100) doubted the possibility of a close affinity between nototylines and paussines as suggested by the above-mentioned authors. Deuve (1994b) published a detailed description of the structural character states of the species and suggested, but with some doubt, a sister-group relationship between nototylines and paussines. He noted several synapomorphies between the two groups including the compressed protibia, the tergite IX which is differentiated into a thin transverse arch, the reduced and lateral position of the laterotergite IX, and the diffuse dorsal pubescence.

Subfamily Paussinae. There is little doubt that this subfamily constitutes a monophyletic lineage. The known larvae share a unique transformation of the abdomen in which the epipleurites of the 9th segment are greatly enlarged and fused with the tergum of the 8th segment to form a plate, displacing the urogomphi and the 10th segment in a vertical plane (Bousquet 1986). The relationship of the subfamily is highly debated but it could be closely related to brachinines. Adults of both groups possess a two-chambered pygidial gland which produces a quinonoid secretion by mixing hydroquinones and hydrogen peroxide from the inner chamber with enzymes produced in the outer chamber (Schildknecht and Holoubek 1961); the secretion is discharged at temperatures of 55-100°C (Aneshansley et al. 1969; Aneshansley et al. 1983). The structure of the pygidial glands and the chemistry of the secretions are unique among beetles. However, based on structural dissimilarities, several authors, including Ball and McCleve (1990), Beutel (1992b), and Geiselhardt et al. (2007), believed that the similarities in the pygidial gland structures and secretions between the two groups are convergent.

Erwin and Stork (1985: 445) concluded that paussines and brachinines are closely related and form the sister-group to a large clade comprising {Elaphrini + Migadopini + Siagonini + Promecognathini + Hiletini + Pseudomorphini + Cnemacanthini (= Cnemalobini) + Scaritini} based on a suite of character states associated with tarsal claws. Deuve (1988), working on the structures of the last abdominal segments of adults, supported the view of a close relationship between paussines and brachinines. However, alternate placements of the paussines have been proposed. Jeannel (1941b: 89) placed trachypachids, gehringiines, and paussines in his Isochaeta based on the apical position of both protibial spurs. Kryzhanovskij (1976a: 87), followed by Lawrence and Newton (1995), included the Cicindini and Nototylini within the Paussinae, implying a close relationship between these three elements. Beutel (1995) suggested a close affinity between paussines and gehringiines. Liebherr and Will’s (1998) preferred cladogram based on 20 characters of the female ovipositors and reproductive tract placed paussines as the sister-group to the remaining Geadephaga (excluding trachypachids). An interesting observation is that of Vigna Taglianti and Rossi (1998: 516) who noted the similarity between the laboulbeniale parasitic species found on the brachinine Pheropsophus Solier and paussine Pachyteles Perty. They added that paussines and brachinines “might be more closely related than suggested by morphological data, thus supporting the result of recent biochemical studies on explosive secretions of members of these groups.”

Members of this subfamily are currently arrayed in five family-group taxa which have been ranked differently during the past few decades. In this catalogue, they are ranked as tribes. All five are probably monophyletic except for the Ozaenini which is likely paraphyletic. The phylogenetic relationships among extinct and extant genera have been expressed in a cladogram based on adult and larval characters by Geiselhardt et al. (2007: Fig. 1).

Tribe Metriini. This group includes two genera: Metrius Eschscholtz, with two species in western North America, and Sinometrius Wrase and Schmidt with a single species recently found in Hubei province in China. This tribe is usually listed as the sister-group to the remaining paussines because of the lack of the apico-lateral fold on each elytron (flange of Coanda of Stork 1985) characteristic of the remaining paussines. This fold, located at the opening of the defence gland, is apparently used to deflect discharges of secretions from the defence glands as showed by Eisner and Aneshansley (1982) for the Neotropical genus Goniotropis Gray. However, Vigna Taglianti et al. (1998: 292), based on a set of 20 larval characters, considered {Metriini + Ozaenini} as the sister-group to Paussini, suggesting that the elytral fold was secondarily lost in metriines or that the fold evolved twice in the subfamily.

Tribe Mystropomini. This tribe includes only the genus Mystropomus Chaudoir, with two Australian species. It is probably the most primitive extant genus of the subfamily excluding metriines. Adult ozaenines, protopaussines, and paussines (sensu stricto) are synapomorphic in having the elytral fold short, the pterothorax and abdomen parallel-sided and the epimera and anepisterna largely covered by the elytral epipleura (Beutel 1992c: 56). In adults of Mystropomus the elytral fold is markedly long and extends over the apical half of the elytron (Jeannel 1946: 47).

Tribe Ozaenini. This group of about 160 species is mainly represented in the tropics; only a few species enter the southern parts of the Northern Hemisphere in Japan, China, Taiwan, and southern United States. Ozaenines differ from protopaussines and paussines by having the mouthparts not modified, and from paussines also in having all 11 antennomeres normally developed. Several authors (e.g., Ball and McCleve 1990; Nagel 1997: 356; Di Giulio and Moore 2004) believed that ozaenines are paraphyletic in regard to the remaining Paussinae (excluding mystropomines and metriines). Beutel (1992b; 1995) and Di Giulio et al. (2003) proposed that the ozaenine genus Physea Brullé is the sister-group to {protopaussines + paussines} while Ball (in Nagel 1997: 356) regarded Ozaena Olivier as the best candidate based on the enlarged first antennomere and the reduced antennal cleaner of the protibia.

Tribe Protopaussini. This tribe includes eight extant Asian species placed in the genus Protopaussus Gestro. Some authors (e.g., Basilewsky 1953a: 23, 1962a: 6-9; Nagel 1987: 27) associated protopaussines with ozaenines based on the presence of 11 antennomeres in both groups but most have associated them with paussines. Nagel (1997: 348, 356) did not find any derived character states shared between protopaussines and ozaenines but noted that the small lacinia lacking the dense brushlike pilosity, typical of other carabids, is a putative synapomorphy for protopaussines and paussines (sensu stricto). From a zoogeographic point of view, it is interesting to note that a Tertiary fossil species of Protopaussus has been described from Dominican amber (Nagel 1997).

Tribe Paussini. This group, also known under the vernacular name “ant nest beetles, ” currently includes about 565 myrmecophilous species arrayed in this work in seven subtribes: Carabidomemnina for the genera Eohomopterus Wasmann (two Neotropical species) and Carabidomemnus Kolbe (27 African species); Arthropterina for the Australian genera Megalopaussus Lea (one species) and Arthropterus Macleay (about 65 species); Cerapterina for the genera Mesarthropterus Wasmann (one species in Ethiopia) and Cerapterus Swederus (32 species in the Afrotropical and Oriental Regions with two species extending into the Himalayas); Pentaplatarthrina for the genera Hexaplatarthrus Jeannel (one Madagascan species) and Pentaplatarthrus Westwood (eight Afrotropical species); Homopterina for the genus Homopterus Westwood (12 Neotropical species); Heteropaussina for the genus Heteropaussus Thomson (about 25 species in the Afrotropical and Oriental Regions); and Paussina for the remaining 12 genera (about 385 species). Luna de Carvalho (1989: 361) used a different approach and recognized three tribes among his extant Paussinae (Paussini in this work): Cerapterini (including carabidomemnines, homopterines, heteropaussines, and arthropterines), Pentaplatarthrini, and Paussini. Within his Paussini, he included the following subtribes: Platyrhopalina for the Asian genera Platyrhopalopsis Desneux (three species), Platyrhopalus Westwood (14 species), Stenorhopalus Wasmann (two species), Lebioderus Westwood (seven species), and Euplatyrhopalus Desneux (six species); Ceratoderina for the genera Paussomorphus Raffray (three Afrotropical species), Melanospilus Westwood (three Oriental species with one species extending into the Himalayas), and Ceratoderus Westwood (seven Asian species); Leleupaussina for the genus Leleupaussus Luna de Carvalho (one Afrotropical species); Hylotorina for the Afrotropical genera Granulopaussus Kolbe (four species), Hylopaussus Luna de Carvalho (two species), and Hylotorus Dalman (six species); and Paussina for numerous genera that some authors sink into one large genus, Paussus Linnaeus (about 330 species in the Old World of which only two, Paussus favieri Fairmaire and Paussus turcicus Frivaldszk von Frivald, reach Europe). Nagel (1987, 1997, as Carabidomemnitae) viewed the Carabidomemnina as the sister-group of the remaining Paussini.

Subfamily Brachininae. There is little doubt that this group, known under the vernacular name “bombardier beetles, ” constitutes a monophyletic lineage. The adults have seven (females) or eight (males) exposed abdominal sterna instead of six as in other carabids. Such modification provides a greater abdominal mobility, allowing a more efficient alignment of the defence spray. However, brachinines do not appear monophyletic in terms of their 18S rDNA (Maddison et al. 1999: 129). The group has a worldwide distribution but is clearly more diverse in the Southern Hemisphere. Most authors recognize two main lineages, ranked here as tribes, among brachinines: Brachinini, represented in most regions of the world including North America, and Crepidogastrini, restricted to southern India and Africa.

For a long time brachinines have been associated with the “Truncatipennes, ” an informal name use to group several tribes whose adults have more or less truncate elytra at the apex. Jeannel (1942, 1949a) included brachinines and pseudomorphines in his Balteifera, implicitly suggesting a close affinity between the two groups. Liebherr and Will (1998: 152-153) placed brachinines with the {Harpalinae + Trechinae + Moriomorphinae}in their study of the female reproductive tract. These authors also alluded to the possibility of a close relationship between brachinines and clivinines. Analysis of molecular data presented by Ribera et al. (2005: 289) indicated a close relationship between brachinines and the subfamily Harpalinae, not with the Paussinae. Maddison et al. (1999: 129) suggested, from 18S r-DNA sequence analyses, an intriguing possibility, that the paussines and brachinines are closely related and that both in turn are related to Harpalinae. In my opinion their hypothesis is credible.

Tribe Crepidogastrini. This tribe is mostly represented in the Afrotropical Region but a few species are found in the Indian subcontinent. It contains the genera Brachynillus Reitter (three species), Crepidogaster Boheman (about 100 species), Crepidogastrillus Basilewsky (one species), Crepidogastrinus Basilewsky (one species), Crepidolomus Basilewsky (two species), and Crepidonellus Basilewsky (five species).

According to Erwin (1970a: 27), adults of crepidogastrines differ from those of brachinines in having the mesepimeron absent or almost so (instead of broad), the adhesive setae on the male protarsi of the “spongy” type (instead of the “seriate” type), the terminal palpomeres swollen and usually securiform (instead of subcylindrical or wedge-shaped), and the gular suture convergent behind (instead of divergent).

Tribe Brachinini. This tribe includes about 540 species of which 50, all belonging to the genus Brachinus Weber, occur in North America. Erwin’s (1970a: 175) study suggested that all New World species of Brachinus, along with a relict species found in the Himalayas, form a clade for which he proposed the subgeneric name Neobrachinus. He also postulated that the subgenus Cnecostolus Reitter, endemic to the Palaearctic Region, was the sister-group to Neobrachinus. Erwin (1970a: 28) arrayed the brachinine genera into four subtribes: Aptinina, Brachinina, Mastacina, and Pheropsophina. In his cladistic analysis (Erwin 1970a: Fig. 451), masticines were positioned as the sister-group to pheropsophines and the two form the sister-group to {aptinines + brachinines}.

Unlike most carabid larvae, those of brachinines are ectoparasites and feed on carabid and water beetle pupae.

Subfamily Harpalinae. Harpalinae is the largest subfamily of Carabidae and the one usually placed at the end of the carabid classification. Molecular data analyses (Maddison et al. 1999; Ober 2002; Ribera et al. 2005) suggest that the subfamily is monophyletic.

In this catalogue, members of Harpalinae are arrayed conveniently in two supertribes: Pterostichitae and Harpalitae. Adults of the vast majority of Pterostichitae, which includes the tribes Morionini, Abacetini, Pterostichini, Zabrini, Oodini, Panagaeini, and Chlaeniini in North America, have crossed epipleura and most secrete something else than formic acid as major constituent of the pygidial glands. Adults of Harpalitae have non-crossed epipleura and, except in the sole species of Pentagonicini studied, secrete formic acid as major constituent of their pygidial glands as far as known. The absence of a crossed epipleuron could be an evolutionary feature providing greater flexibility to aim the powerful formic acid secretion of the pygidial glands. The presence of a transverse membranous band on the stipes of larvae prompted Arndt (1998: 184) to suggest that the tribes Licinini and Harpalini, herein included in the Harpalitae, were closely related to members of Pterostichitae.

Tribe Morionini. This relatively well-defined, likely monophyletic tribe is represented in all zoogeographical regions of the world but is more diverse in the tropics than in temperate areas. Its relationships have been debated. Indeed, some larval character states suggest that morionines could be related to scaritines while some adult character states suggest they may be related to pterostichines. Bousquet (2001) discussed the larval character states of morionines in detail and concluded that they do not yield evidence to favor one hypothesis over the other. However, when features of the adults are also taken into account, there is little doubt that morionines are more closely related to pterostichines than to scaritines. Recently Will (2004: 218), following Liebherr and Will (1998: 156), found three “unambiguously optimized and unreversed synapomorphies” suggesting that cnemalobines and morionines are sister-groups. A review and cladistic analysis of the morionine genus-group taxa have been published recently (Will 2004).

Moore (1965: 5) included the Australian genus Catadromus Macleay (seven species) in the tribe Morionini but his view has not been retained by subsequent authors.

Tribe Cnemalobini. This tribe includes only the Neotropical genus Cnemalobus Guérin-Méneville (32 species in Chile, Argentina, and Uruguay). Jeannel (1941b: 286) stated that the genus should be placed near the perigonines and Reichardt (1977: 416) followed his suggestion. Erwin (1985: 467) associated cnemalobines (as Cnemacanthini) with scaritines and clivinines. Arndt (1993: 40) suggested that the tribes Cnemalobini and Harpalini form a clade based on larval characteristics. Roig-Juñent (1993: 12) suggested, from a preliminary analysis, that cnemalobines and zabrines are sister-groups and the two groups are closely related to morionines. Other cladistic analyses (Liebherr and Will 1998: 156; Will 2004: 217) placed morionines as the sister-group to cnemalobines. Molecular data (18S rDNA) analyses (Maddison et al. 1999: 129) did not endorse placement of cnemalobines with Scaritinae but supported an association with the subfamily Harpalinae.

Tribe Microcheilini. This tribe includes a single genus, Microcheila Brullé, with two Madagascan species. Besides their relatively aberrant facies, adults of this group possess a number of character states unusual for pterostichines. The penultimate labial palpomere has more than two setae, each sternum possesses a transverse row of setae, the protibia has a latero-apical dentiform protuberance, all tarsomeres are densely pubescent beneath, and the first four protarsomeres of the male have adhesive setae (Jeannel 1948a: 616). The elytral plica is well developed as in members of Pterostichini. The group was included, along with morionines, chaetodactylines, and pterostichines (including sphodrines and platynines), in Jeannel’s (1948a: 380) family Pterostichidae.

Tribe Chaetodactylini. This group includes a single genus, Chaetodactyla Tschitschérine with 20 species endemic to Madagascar. The species superficially resemble several pterostichine taxa but the male protarsomeres are not expanded and have no adhesive setae (Jeannel 1948a: 619). The group was associated with morionines, metiines, zabrines, microcheilines, and pterostichines (including sphodrines and platynines) in Jeannel’s (1942: 734-735) family Pterostichidae.

Alluaud (1935: 28) reported that one of his colleagues rearing pupae of various insect groups for parasitic Hymenoptera discovered 14 adults of Chaetodactyla emerging from pupal chambers of two cetonid species. Jeannel (1948a: 620) postulated that Chaetodactyla females probably lay their eggs on the cetoniid larvae and that the carabid larvae remain inside the cetonid pupae, eventually feeding upon them.

Tribe Cratocerini. This tribe includes the genera Cratocerus Dejean with two Neotropical species and Brachidius Chaudoir with one australo-oriental species. Chaudoir (1873a) also listed Basoleia Westwood (= Catapiesis Solier) in this tribe and Lorenz (2005: 248) also included the genus Oxyglychus Straneo, with one Japanese species, previously included within the caelostomines (= drimostomatines). Cratocerines have been little studied and their taxonomic position is not well established. They are usually placed within the Pterostichini (e.g., Reichardt 1977: 407). Lorenz (2005: 248-252) combined cratocerines with catapieseines and drimostomatines in his subfamily Pterostichinae.

Tribe Abacetini. This tribe is proposed here to include the abacetines proper, the loxandrines, and the celioscheseines based on a preliminary cladistic analysis conducted by Will (2000) suggesting that these three groups are closely related. Van Emden (1949) and Arndt (1988) had already drawn attention to the fact that some putative apomorphic character states were shared by abacetines (with more than 95% of the species endemic to the Old World) and loxandrines (with more than 95% of the species restricted to the New World). As defined here, this tribe, as well as all three groups included in it, is inadequately characterized except for some abacetine genera which have an asymmetrical insertion of the second antennomere in the adults, and some loxandrine genera which have the first three protarsomeres of the males obliquely expanded. Monophyly of this tribe has not yet been demonstrated.

Tribe Pterostichini. This highly diverse tribe is represented in all continents, except Antarctica, and the species are found from the arctic regions to the tropics. There are no structural features yet discovered to suggest that the tribe, as currently conceived, forms a clade and there is little doubt, as suggested by Ball (1979: 102), that it represents a grade.

A number of putative clades have been recognized within the pterostichines and some of them have received formal scientific names. These include, among others, the euchroines with the genera Bothynoproctus Tschitschérine (one Neotropical species), Euchroa Brullé (38 Neotropical species), Lobobrachus Sharp (two Neotropical species), Setalis Laporte (three Australian species) and, according to Will (2000: 64), Microcephalus Dejean (15 Neotropical species); the Northern Hemisphere myadines with the genus-group taxa Aristochroa Tschitschérine (18 Asian species), Myas Sturm (with about 30 species in North America and Asia placed in the subgenus Trigonognatha Motschulsky and one European species), Steropanus Fairmaire (11 Asian species, some of them endemic to the Oriental Region), and Xenion Tschitschérine (one European species) to which Stereocerus Kirby (two Holarctic species) is probably closely related (Bousquet 1999: 85); the trigonotomines (including deliniines) with the genera Delinius Westwood (three Australian species), Leiradira Laporte (12 Australian species), Lesticus Dejean (about 100 Asio-Australian species), Trigonotoma Dejean (about 55 Asian species), and Euryaptus Bates (six Asian species), Pareuryaptus Dubault, Lassalle and Roux (17 Asian species); the Australian darodiliines (including cratogastrines) with the genera Loxogenius Sloane (one species), Liopasa Tschitschérine (one species), Cratogaster Blanchard (five species), and Darodilia Laporte (ten species); the New Caledonian abacomorphines with the genera Abacoleptus Fauvel (three species), Abacomorphus Chaudoir (two species), Platysmodes Fauvel (one species), and Setalidius Chaudoir (two species); the molopines with the North American genus Cyclotrachelus Chaudoir (45 species) and the western Palaearctic genera Abax Bonelli (18 species), Henrotiochoromus Busulini (one species), Molopidius Jeannel (one species), Molops Bonelli (40 species), Oscadytes Lagar Mascaro (one species), Percus Bonelli (19 species), Speomolops Patrizi (one species), Stenochoromus Miller (one species), Styracoderus Chaudoir (three species), Tanythrix Schaum (three species), Typhlochoromus Moczarski (two species), and Zariquieya Jeannel (one species) to which Jeannel (1948a: 450-451) added several Madagascan genera (Abacodes Jeannel, Eucamptognathus Chaudoir, Eudromus Klug, Eurypercus Jeannel, and Molopinus Jeannel); and the poecilines as defined by Jeannel (1942: 738) with the genera Stomis Clairville, Pedius Motschulsky, Argutor Dejean, Orthomus Chaudoir, Poecilus Bonelli, Phonias des Gozis, Bothriopterus Chaudoir, and Melanius Bonelli. Some of these groups, such as the poecilines, are probably polyphyletic.

Tribe Zabrini. Zabrines are most diversified in the Palaearctic and Nearctic Regions but are also represented in the mountains of the northern Neotropical, northern Oriental, and eastern Afrotropical Regions. Some authors have recognized several, more or less clearly defined genera in this tribe, others only two, Amara Bonelli and Zabrus Clairville, each with many subgenera. Adults of zabrines are structurally most similar to members of Pterostichini and probably represent a clade within the Pterostichini as presently conceived.

Tribe Metiini. This tribe includes about 75 species restricted to the southern part of South America, predominantly in Chile and extending north to Peru and east to southern Brazil, Uruguay, and Argentina. These species are arrayed in the following genera: Kuschelinus Straneo (one species), Metius Curtis (about 60 species), Abropus Waterhouse (one species), Antarctiola Straneo (four species) and, according to Will (2000: 60), Feroniola Tschitschérine (nine species). Metiines are often included within the Pterostichini.

This tribe has been known in the past under the name Antarctiini. However, because its type genus Antarctia Dejean is a junior homonym, the family-group name Antarctiini is permanently invalid (ICZN 1999: Article 39).

Tribe Drimostomatini (including cyrtolaines). The association of the Eastern Hemisphere drimostomatines (also known under the name caelostomines) with the Western Hemisphere cyrtolaines (Cyrtolaus Bates with 11 Middle American species and Barylaus Liebherr with two species in the West Indies) was proposed by Liebherr (1986) and supported by Will’s (2000) preliminary cladistic analysis. The main characteristic of this group is the inverted aedeagus. However, this modification is absent in some groups (e.g., Diceromerus Chaudoir) traditionally placed within the drimostomatines and consequently monophyly of this tribe is uncertain. The drimostomatines include about 290 species arrayed in 29 genera (Lorenz 2005: 248-252, as Drimostomatina). The most speciose genera are Caelostomus Macleay (about 160 species, of which one is adventive in the West Indies), Trichillinus Straneo (21 species), Platyxythrius Lorenz (20 species), and Strigomerus Chaudoir (18 species).

The name Caelostomini, proposed by Burgeon (1935: 194), is often used for this tribe but Drimostomatini, established by Chaudoir (1872c: 283), is older and has priority. Drimostoma Dejean is usually treated as a junior synonym of Caelostomus Macleay but the family-group name Caelostomini was not proposed because of the synonymy of the type genus. Therefore, Article 40.2 of the ICZN (1999) does not apply in this case.

Tribe Chaetogenyini. This South American tribe includes five species of the genus Chaetogenys van Emden arrayed in two subgenera: Chaetogenys s.str. and Camptotoma Reiche. The group has been ranked as a subtribe of Pterostichini by some authors, including van Emden (1958), Straneo (1977), and Reichardt (1977: 408). However, the adhesive setae on the male protarsi are of the “spongy” type (Reichardt 1977: 408), not of the “seriate” type as in other pterostichines. Erwin (1985: 468) associated chaetogenyines with cuneipectines, chlaeniines, oodines, and licinines.

Tribe Dercylini. The 35 species of this exclusively Neotropical tribe are currently arrayed in one genus (Dercylus Laporte) with four subgenera (Moret and Bousquet 1995: 759): Asporina Laporte (two species), Dercylus s.str., with Dercylodes Chaudoir and Pterodercylus Kuntzen as synonyms (12 species), Eurydercylus Moret and Bousquet (seven species), and Licinodercylus Kuntzen, with Physomerus Chaudoir (a junior homonym) as synonym (14 species). Chaudoir (1883), Reichardt (1977), and Ball (1979: 102) suggested that dercylines were closely related to oodines. Moret and Bousquet (1995: 759) stated that the character states of the adult and of the putative larva studied indicate that dercylines are more closely related to oodines and chlaeniines than to pterostichines. Bousquet (1996a: 449) commented that dercylines were closely related to {oodines + panagaeines + chlaeniines} but that the nature of the relationship remained to be ascertained. Jeannel (1948a: 626) related dercylines to melanchitonines and Kryzhanovskij (1976a: 89) to pterostichines, microcheilines, chaetodactylines, platynines, zabrines, and cuneipectines without mentioning any character state that would justify such grouping. The adhesive setae on the male protarsi are of the “spongy” type as in chaetogenyines, oodines, and chlaeniines.

Jeannel (1948a: 627) indicated that the genus Dercylinus (one North American species), of which he had seen no specimen, probably belongs to dercylines and Lorenz (2005: 327) listed the genus, along with Evolenes (one North American species), in the subtribe Dercylina. However, these two genera are typical oodines (see Bousquet 1996a).

Tribe Melanchitonini. This tribe currently includes three genera, Melanchiton Andrewes (a replacement name for Melanodes Chaudoir), Melanchrous Andrewes (a replacement name for Patellus Chaudoir), and Dicaelindus Macleay. The lineage contains about 70 Old World species. As for many other groups, relationships of melanchitonines are unclear. Chaudoir (1883) included Melanchiton and Melanchrous within the tribe Oodini, likely because of the similar adhesive setae on the male protarsi. Subsequently, the two genera have been placed by some authors within the Pterostichini. Jeannel (1948a: 626) included them with dercylines in his family Dercylidae but offered no pertinent evidence to suggest that the group is monophyletic.

Straneo (1950: 65) first included the genus Dicaelindus, previously placed in the Pterostichini, in this tribe. Adults of Dicaelindus are rather similar phenetically to those of Melanchiton, but the male protarsi are not dilated and lack adhesive setae. Monophyly of this tribe has not yet been demonstrated.

Tribe Oodini. Members of Oodini sensu stricto share several apomorphic character states in the adult stage (Bousquet 1996a: 448) suggesting the tribe is monophyletic. Several authors have included or associated oodines with chlaeniines but the pygidial gland components suggest rather that panagaeines and chlaeniines are more closely related to each other than to oodines (Bousquet 1987b). Oodines, panagaeines, and chlaeniines possibly constitute a clade since the adults (except in some chlaeniines) have the metepisterna coadunate with the elytral epipleura, a synapomorphic condition that has probably been secondarily lost in some chlaeniine lineages.

Some groups, such as dercylines, melanchitonines, and geobaenines, are sometimes included within the Oodini as distinct subtribes. However, there is little evidence that they are indeed closely related to oodines and in my opinion they should be treated as distinct tribes.

This tribe is represented in all zoogeographical regions of the world and includes about 295 species in 32 genera. Jeannel (1949a: 829) recognized three family-group taxa within the oodines: sphoerodines represented in the Afrotropical Region, oodines (sensu stricto) represented in all zoogeographical regions, and thryptocerines represented in the Afrotropical Region.

Tribe Peleciini. Relationships of peleciines are unclear. The group has been associated with panagaeines by Kryzhanovskij (1976a: 89), Ball (1979), and Erwin (1985: 468) and included in the superfamily Odacanthomorphi, along with odacanthines, perigonines, lachnophorines, and ctenodactylines, by Jeannel (1948a: 376). Many apomorphic features, including some details of the chaetotaxy, structure of the mouthparts and thorax, and marked similarity in their specialized way of attacking millipedes, suggest that peleciines could be closely related to promecognathines. However, Straneo and Ball (1989) regarded these similarities as evolutionary convergence, not phylogenetic affinity. Larvae of Eripus oaxacanus Straneo and Ball, the only peleciine species known in its larval stage, are similar in some structural features to larvae of Brachinini and Pseudomorphini but Liebherr and Ball (1990) concluded that these similarities were an example of convergence due to a similar parasitic lifestyle. Arndt (1993: 36), based on larval features, suggested that peleciines, panagaeines, licinines, oodines, and chlaeniines form a clade. Liebherr and Will (1998: 156-157) noted from their analysis of the female reproductive tract that placement of peleciines as a basal group of pterostichine stock was firmly supported.

Peleciines are restricted to the Southern Hemisphere. Straneo and Ball (1989) recognized two subtribes: Agonicina for the genera Pseudagonica Moore and Agonica Sloane of Tasmania and adjacent southeastern Australia, and Peleciina (including disphericines) for the remaining genera which are represented in the Afrotropical, Oriental, and Neotropical Regions. Vigna Taglianti and Rossi (1998: 515) noted that the laboulbeniale parasitic species found in Agonica and in the moriomorphine genera Pterogmus Sloane, Theprisa Moore, and Sitaphe Moore were very similar and alluded to the possibility of a close relationship between agonicines and moriomorphines.

Tribe Brachygnathini. This tribe contains only the Neotropical genus Brachygnathus Perty (seven species). Relationships of the genus are uncertain. Jeannel (1949a: 849) associated it with the genus Microcephalus Dejean (as Tichonia Semenov), under the subfamily name Tichoniitae, and placed it in his family Panagaeidae. Reichardt (1977: 404) noted that inclusion of Brachygnathus in the tribe Panagaeini was doubtful and that the adults show some similarities to those of peleciines.

Tribe Bascanini. This tribe contains a single genus, Bascanus Péringuey (including Bascanidius Péringuey), with a few species in eastern and southern Africa. Van Emden (1936a), Basilewsky (1953a: 164-165), and Erwin (1979) suggested that bascanines are closely related to panagaeines. Csiki (1933a: 1651) associated the genus with Melaenus Dejean.

Tribe Panagaeini. This moderately diverse group occurs in all continents except Antarctica but is much more diverse in the tropics than in temperate regions. Panagaeines, at least those that have been analysed, secrete phenol through their pygidial glands (see Schildknecht et al. 1968; Kanehisa and Murase 1977; Moore 1979). This compound is also found, as far as known, only in some chlaeniines, which suggests that panagaeines are probably most closely related to chlaeniines. On the other hand, several authors, including Kryzhanovskij (1976a: 89), consider peleciines as the group most closely related to panagaeines.

Jeannel (1949a: 849) associated the genus Microcephala Dejean (as Tichonia Semenov) with Panagaeini but most authors, including Reichardt (1977: 407), regard it as a member of Pterostichini.

Tribe Chlaeniini. Chlaeniines are found in all zoogeographical regions of the world but are more diverse, both in terms of lineages and species, in the Afrotropical and Oriental Regions than anywhere else. Jeannel (1949a: 776) recognized six tribes among chlaeniines and Basilewsky and Grundmann (1955) ten tribes and two subfamilies. However, following Ball (1960b) and Lindroth (1969a), all the species are grouped in a single tribe in this catalogue. Several authors have suggested a close relationship between chlaeniines, panagaeines, and oodines.

Two major groups among Chlaenius species could be distinguished based on defensive secretions of the pygidial glands (see Schildknecht et al. 1968; Kanehisa and Murase 1977; Moore 1979; Balestrazzi et al. 1985): one secretes phenol, like panagaeines; the other one quinone. In the first group, the secretory lobes of the pygidial glands are elongate, in the second one they are shorter and thicker (Kanehisa and Shiraga 1978). I believe these two groups should be recognized either as genera or subtribes. However, owing to the lack of information on the pygidial glands and their secretions for many chlaeniine lineages, such action is futile at this time.

This tribe includes almost a thousand species worldwide arranged in 18 genera and two subtribes. The 51 North American species are assigned to the genus Chlaenius Bonelli and arrayed in ten subgenera of which five, Pseudanomoglossus Bell (one species), Anomoglossus Chaudoir (three species), Callistometus Grundmann (one species), Brachylobus Chaudoir (one species), and Randallius n.subg. (one species), are North American endemics.

Tribe Cuneipectini. This tribe includes one genus, Cuneipectus Sloane, with two flightless species in western Australia. Members of this group have rarely been collected and very little is known about their way of life. Kryzhanovskij (1976a: 89) listed cuneipectines in his supertribe Pterostichitae along with dercylines, zabrines, platynines, chaetodactylines, microcheilines, and pterostichines. Erwin (1985: 468) associated them with chaetogenyines, chlaeniines, oodines, and licinines in his supertribe Callistitae (= Chlaeniitae). Moore et al. (1987: 215) included them with morionines, pterostichines, abacetines, geobaenines, drimostomatines, and platynines in their Pterostichitae.

Tribe Orthogoniini. This group includes six genera represented in Asia and Africa only: Orthogonius Macleay (about 240 species), Neoorthogonius Tian and Deuve (one species), Hexachaetus Chaudoir (nine species), Actenoncus Chaudoir (four species), Anoncopeucus Chaudoir (two species), and Nepalorthogonius Habu (one species). Relationships of the tribe remain unresolved and problematic. Jeannel (1948a: 377) indicated that orthogoniines and licinines are closely related based on the shape of the frontale on the cephalic capsule of the larvae. Basilewsky (1953a: 180) associated them with glyptines, Kryzhanovskij (1976a: 90) with lebiines, anthiines, helluonines, physocrotaphines, zuphiines, galeritines, and dryptines, and Erwin (1985: 468) with idiomorphines, catapieseines, and amorphomerines. Ober and Maddison (2008: 18) found strong support in their phylogenetic analyses based on molecular data sequences for a clade comprising orthogoniines, graphipterines, and pseudomorphines. The genus Glyptus Brullé has been included by some authors within the tribe Orthogoniini, but both Jeannel (1948a) and Erwin (1985) believe that Glyptus and Orthogoniini are not closely related. Members of this tribe are termitophilous.

Tribe Idiomorphini. This tribe currently includes the genera Idiomorphus Chaudoir (three Indian species), Perochnoristhus Basilewsky (one species in Namibia), Rathymus Dejean (three Afrotropical species), and Strigia Brullé (three Oriental species) arrayed in two subtribes, Perochnoristhina for the genus Perochnoristhus and Idiomorphina for the remaining genera (Lorenz 2005: 391). Erwin (1984b: 378) also included the genus Glyptus in this tribe. Crowson (1980) stated that the genus Perochnoristhus could be closely related to broscines and apotomines.

Tribe Glyptini. Glyptini consists of two Afrotropical genera: Neoglyptus Basilewsky with six species and Glyptus Brullé with two species. Few authors agree on the systematic position of the group. Jeannel (1948a: 377) associated them with chlaeniines, Basilewsky (1953a: 180) with orthogoniines, and Erwin (1984b: 378) with idiomorphines. Both Chaudoir (1850a) and Lacordaire (1854) stated that glyptines were closely related to the genus Idiomorphus Chaudoir.

Tribe Amorphomerini. This group includes a single genus, Amorphomerus Sloane, represented by a few species in eastern Africa and Madagascar. Jeannel (1948a: 376) associated amorphomerines with pterostichines (sensu lato, including platynines), dercylines, and harpalines in his superfamily Harpalomorphi, characterized by having the mesotibiae spinose and the median lobes of the aedeagi more or less bent, with the basal bulbs well developed. He also stated (Jeannel 1948a: 731) that the tribe was more closely related to harpalines than to any other Conchifera groups. Kryzhanovskij (1976a: 89) associated amorphomerines with harpalines, cnemalobines (as Cnemacanthini), and agonicines (currently included in the Peleciini) and Erwin (1985: 468) associated them with idiomorphines, orthogoniines, and catapieseines. The tribe was listed as part of the tribe Lebiini by Erwin (1979).

Tribe Licinini. A clearly defined, likely monophyletic group with representatives in all zoogeographical regions of the world. Jeannel (1948a: 377) associated licinines with pentagonicines, orthogoniines, panagaeines, chlaeniines (including oodines), and glyptines, Kryzhanovskij (1976a: 89) with oodines and chlaeniines, and Erwin (1991a: 10) with oodines, chaetogenyines, chlaeniines, and cuneipectines. Ball (1992a) considered the tribe to be the sister-group to {Oodini + Chlaeniini + Panagaeini} and Ball and Bousquet (2000: 100) noted that members of the four tribes show similarities in structure of the male protarsi, genitalia, and larvae. Beutel (1992d) reported several putative synapomorphies in larval head structures between Licinini and Panagaeini, and Arndt (1993: 37) noted several synapomorphies in larvae of licinines, panagaeines, and peleciines. However, contrary to the oodine-chlaeniine-panagaeine complex, licinines have simple (i.e., non-crossed) epipleura and secrete formic acid as the major constituent of their defensive glands like harpalines and Truncatipennes members. Also Ober and Maddison (2008: 19) found no close relationship between licinines and the chlaeniine-oodine-panagaeine complex based on their analyses derived from molecular data sequences. Recently Liebherr and Will (1998: 144) suggested that licinines, orthogonines, panagaeines, melanchitonines, graphipterines, and loxandrines form a clade based on the presence of a villous canal extended forward on the common oviduct.

The 235 or so species are arrayed in 23 genera distributed among four subtribes following Ball (1992a).

Tribe Harpalini. This is one of the largest and most diversified carabid tribes. Although its limits are fairly stable, there is as yet no strong evidence to substantiate that the tribe is monophyletic. Based on a study of the world fauna, Noonan (1976) recognized four subtribes among harpalines: Anisodactylina, Pelmatellina, Stenolophina (including polpochilines and pachytrachelines), and Harpalina which he divided into eight genus-groups, namely Harpali, Selenophori, Bradybaeni, Acinopi, Bleusei, Dapti, Amblystomi, and Ditomi. Based on a parsimony analysis of molecular sequence data, Martínez-Navarro et al. (2005) concluded that the subtribe Harpalina was polyphyletic, that daptines were related to stenolophines, not to Harpalina, that the Selenophori group was polyphyletic and not related to Harpalina but perhaps to anisodactylines, that the Amblystomi group may be related to stenolophines instead of Harpalina, and that the subtribe Pelmatellina was related to stenolophines (see also Martínez-Navarro et al. 2003) and that the latter could be paraphyletic in regard to the former. They also advocated raising selenophorines, ditomines, and amblystomines to subtribe level.

Relationships of harpalines to other carabid groups are not well established. In the course of his work on the French fauna, Jeannel (1942: 575) associated harpalines with perigonines, anchonoderines, lachnophorines, omphreines, pterostichines (including platynines), zabrines, chaetodactylines, morionines, and metiines in his superfamily Harpalomorphi. Later, working on the Madagascan fauna, Jeannel (1948a: 376) united the harpalines with amorphomerines, dercylines, melanchitonines, pterostichines (including platynines), morionines, microcheilines, and chaetodactylines. Kryzhanovskij (1976a: 89) listed harpalines with amorphomerines, cnemalobines, and agonicines (currently included in Peleciini) in his supertribe Harpalitae. Based on the presence of a membranous transverse band on the stipes lateroventrally in larvae, Arndt (1998: 184) associated harpalines with morionines, pterostichines, zabrines, panagaeines, peleciines, chlaeniines, oodines, licinines, and cnemalobines. In a cladistic analysis conducted by Roig-Juñent and Cicchino (2001: Fig. 1), this tribe was positioned as the sister-group to {Platynini + Sphodrini}. Ruiz et al. (2008) indicated that, based on their molecular data sequence analyses, the tribe Harpalini was the sister-group to {Sphodrini + Platynini + Pterostichini + Zabrini}.

Tribe Geobaenini. The Geobaenini includes a single genus, Geobaenus Dejean, with four flightless species: three occur in South Africa, one in Australia. The group was first included within the tribe Harpalini and associated subsequently with pterostichines. Basilewsky (1949), because of similarity in the adhesive setae on the male protarsi, suggested that geobaenines could be closely related to melanchitonines, although later (1950, 1953, 1985) he associated the genus with platynines (as Anchomeninae or Platyninae). Liebherr and Will (1998: 144) in their study of the female reproductive tract found an “uncontested synapomorphy” uniting geobaenines with lachnophorines, odacanthines (including pentagonicines), and pseudomorphines. In these taxa, the spermathecal duct is joined to the common oviduct by an elongate sclerite.

Tribe Omphreini. This tribe includes a single genus, Omphreus Dejean (18 species), which is endemic to the Balkan Peninsula and Asia Minor. Omphreines have been included within the tribe Platynini by most authors but Jeannel (1942: 577), followed by Kryzhanovskij (1976a: 89), associated them with perigonines, anchonoderines (including atranines), and lachnophorines.

Tribe Sphodrini. Members of this group have been traditionally included within the Platynini. However, in recent decades numerous taxonomists dealing with the Palaearctic fauna, where this group is by far more diversified than anywhere else, rank this complex as a distinct tribe. Based on morphological characters, there seems to be little doubt that the two groups are closely related. However, from molecular data sequence analyses conducted by Ruiz et al. (2008), this relationship did not receive “the expected strong support, though it can not be completely dismissed.” The Sphodrini include about 825 species, arranged in about 40 genera, and are grouped into the following six subtribes: Atranopsina (about 100 species), Calathina (about 185 species), Dolichina (17 species), Pristosiina (about 65 species), Synuchina (almost 100 species), and Sphodrina (about 360 species). Based on Casale’s (1988: 130) cladogram, Dolichina and Synuchina are sister-groups, and Sphodrina, Calathina, and Pristosiina form a clade with Pristosiina the sister-group to the other two; the position of Atranopsina is ambiguous. From the molecular data sequence analyses conducted by Ruiz et al. (2008), only the position of the subtribe Atranopsina as the sister-group to all other subtribes was well supported.

Tribe Platynini. This is a large, complex, and worldwide group which is more diverse in the tropics than in temperate regions. There are no synapomorphies, in either adult or larval structures, yet discovered to suggest that the tribe represents a monophyletic lineage. Platynines are combined by various authors with pterostichines based on phenetic similarity between the two groups. I believe the two groups are not closely related because of the differences in elytral epipleuron configurations and pygidial gland structures and secretions. Basilewsky (1985, as Platyninae) gave an excellent introduction to the systematics of the group.

Relationships among the North American genus-group taxa have been addressed but are still inadequately understood. According to Liebherr (1991b: 5), Tetraleucus, Anchomenus, Sericoda, and Elliptoleus form a clade characterized by the synapomorphic condition of the female spermatheca having a basal reservoir and a long apical filament. Within this clade Tetraleucus is the sister-group to the remaining taxa. Liebherr and Schmidt’s (2004: 168) parsimony-based cladistic analysis led to the recognition of four subgenera within the genus Agonum forming two clades, {Platynomicrus Casey + Europhilus Chaudoir} and {Agonum s.str. + Agonothorax Motschulsky (= Olisares Motschulsky)}. Liebherr and Schmidt (2004: 153) suggested a sister-group relationship between the genus Agonum and the African taxa described in combination with Agonidium Jeannel and Neobatenus Jeannel as well as several others described under Megalonychus Chaudoir.

Tribe Perigonini. This small tribe is represented by about 115 species arranged in five genera. The place of the tribe within the carabids is unsettled. LeConte and Horn (1883: 35) and Sloane (1923: 248) included it as a separate group within the Platynini; Jeannel (1942: 577) as a distinct subfamily within his family Perigonidae along with anchonoderines, omphreines, and lachnophorines and later (Jeannel 1948a: 376) as a distinct family within his superfamily Odacanthomorphi along with lachnophorines, odacanthines, ctenodactylines, and peleciines. Kryzhanovskij (1976a: 89) followed Jeannel (1942) and combined the tribes Perigonini, Lachnophorini (including anchonoderines), and Omphreini in his supertribe Perigonitae. Erwin (1984b: 375) placed this tribe in his supertribe Lebiitae along with amorphomerines, catapieseines, graphipterines, tetragonoderines, masoreines, pentagonicines, odacanthines, and lebiines. Later (Erwin 1991a: 10) the amorphomerines and catapieseines were excluded from the Lebiitae.

The North American fauna includes two species of the genus Perigona Laporte which contains about 100 species worldwide. One of our species is adventive and the second one is endemic to the eastern part of the continent.

Tribe Ginemini. This tribe includes a single species, Ginema thomasi Ball and Shpeley, known from a single female specimen collected in the departament of Santa Cruz in Bolivia. Ball and Shpeley (2002a: 96) noted some marked similarities between this genus and members of Cyclosomini but still postulated a rather isolated position in the rank of the more derived Harpalinae lineages.

Tribe Enoicini. This tribe includes two South African genera: Enoicus Péringuey with one species and Abacetodes Straneo (= Phimus Péringuey, a preoccupied name) with four species. Basilewsky (1985: 15-16) associated enoicines with platynines, geobaenines, and sphodrines while earlier (Basilewsky 1953a: 61) he included them within the platynines (as Anchomenini).

Tribe Atranini. This tribe contains only two species, both included in the genus Atranus LeConte: one lives in Europe and the Caucasus, the other one in eastern North America. The systematic position of the genus has been debated. Dejean (1828: 122) described the North American species in the genus Anchomenus Bonelli, currently placed within the Platynini. LeConte (1847: 438; 1861a: 28), Seidlitz (1887: 10), and Sloane (1923: 250) associated the genus with chlaeniines, LeConte and Horn (1883: 37), Fauvel (1888: 15), and Jeannel (1942: 582) with anchonoderines, and Ball (1960b: 136), Lindroth (1966: 648), Liebherr (1986: 20), Kryzhanovskij et al. (1995: 118), and several others with platynines. Adults and larvae of Atranus possess several structural features not exhibited in other Platynini. In my opinion the morphological evidence relating Atranus to platynines is weak and for that reason the genus is retained here in its own tribe. Analysis of the pygidial secretions could be useful to indicate if the genus is more closely related to chlaeniines or to the platynine-anchonoderine lineage. Basilewsky (1962b: 155) believed the genus was more closely related to platynines than to any other group suggested to date, but because the adhesive setae on the male protarsi are of the “spongy” type rather than the “seriate” type, he advocated placing it in a distinct subfamily. Phylogenetic relationships as inferred from 28S ribosomal DNA and the wingless gene conducted by Ober and Maddison (2008) placed Atranus as the sister-group to the Platynini.

Tribe Catapieseini. This small Neotropical tribe includes two genera, Catapiesis Brullé with eight species and Homalomorpha Brullé with one species, ranging collectively from southern Mexico to northeastern Argentina. Catapieseines have been placed by some authors (e.g., Reichardt 1977) in the vicinity of the Morionini and Pterostichini. Lorenz (2005: 248) included them within the tribe Cratocerini, in his subfamily Pterostichinae, along with drimostomatines. Ober and Maddison (2008: 16), following Erwin (1984b: 375), placed them in their lebiomorph assemblage pointing out that catapieseines have truncate elytra and specialized eighth abdominal tergite turrets like the remaining lebiomorph taxa. Horn (1881: 163) believed that Catapiesis (as Basoleia) has a close relationship with the Helluonini. Erwin (1985: 468) listed the group within his supertribe Orthogoniitae along with idiomorphines, amorphomerines, and orthogoniines.

Tribe Lachnophorini (including anchonoderines). This small tribe includes about 120 species in ten genera. There is no solid structural or molecular evidence that would suggest this group is monophyletic. Its relationships have been discussed by several authors. Jeannel (1942: 578), followed by Kryzhanovskij (1976a: 89), associated lachnophorines with perigonines, anchonoderines (including Atranus), and omphreines. Liebherr (1988) suggested that lachnophorines derived from a platynine-like ancestor. He also included calophaenines within lachnophorines but Ball and Bousquet (2000: 107), following Erwin (1991b: 44), placed them with ctenodactylines.

The Lachnophorini are represented in the New World, and by one Indo-African species, Selina westermanni Motschulsky. Jeannel (1948a: 744) also considered the genus Amoebea Péringuey (one Afrotropical species) as lachnophorine. However, the name is a junior synonym of Smeringocera Chaudoir (six species) which belongs to the tribe Odacanthini (see Lorenz 2005: 444).

In Liebherr’s (1988) parsimony-based cladistic analysis of the West Indies lachnophorines, the taxa with setose body and setose maxillary palpomeres, represented by the genera Euphorticus, Calybe, and Lachnophorus, constituted a clade.

Tribe Pentagonicini. Few characteristics hold the pentagonicines together and this tribe is possibly polyphyletic. Jeannel (1949a: 767) included pentagonicines in his superfamily Callistomorphi, along with licinines, orthogoniines, panagaeines, chlaeniines, and glyptines, and indicated that pentagonicines are best placed near licinines. Moore (1966a: 162) wrote that larval characters of the pentagonicine Scopodes “suggest a rather close affinity with the Odacanthinae.” Liebherr (1988) included pentagonicines within the Odacanthini based mainly on the structure of the spermatheca, and Ober and Maddison (2008: 17) found support from their molecular data for such a relationship. Erwin (1984b: 375) placed this tribe in his supertribe Lebiitae along with amorphomerines, perigonines, catapieseines, graphipterines, tetragonoderines, masoreines, odacanthines, and lebiines. The author later adopted a similar arrangement (Erwin 1985: 468), with the exclusion of amorphomerines and catapieseines and the inclusion of lachnophorines. It is of interest to note that the sole species of pentagonicines studied (Scopodes boops Erichson) produces a saturated acid as major component (Moore 1979) of the pygidial glands, while members of the so-called Truncatipennes, with which this tribe is usually associated, produce formic acid (see Schildknecht et al. 1968; Moore and Wallbank 1968; Kanehisa and Murase 1977; Moore 1979).

This tribe includes close to 170 species in the world. Only six, all in the genus Pentagonica Schmidt-Göbel (86 species worldwide), are found in North America.

Tribe Odacanthini. The Odacanthini, with about 300 species in 30 genera, constitutes a moderately diverse group represented in all zoogeographical regions. They are more diverse in the tropics than in temperate regions. Several authors believed the group to be closely related to ctenodactylines, and both groups have been combined in a single tribe by some (Csiki 1932b, Liebke 1938, van Emden 1942). Jeannel (1948a: 376) associated odacanthines with perigonines, lachnophorines, ctenodactylines, hexagoniines, and peleciines, and Basilewsky (1962b: 154) with lachnophorines. Liebherr (1988) concluded that odacanthines (including pentagonicines) have a sister-group relationship with lachnophorines (including calophaenines) based on the presence of a bipartite spermatheca. However, phylogenetic analyses based on molecular data sequences presented by Ober and Maddison (2008: 5) did not support odacanthines and lachnophorines as a clade but did support a close relationship between pentagonicines and odacanthines and between calophaenines and lachnophorines. Erwin (1985: 468) placed the tribe Odacanthini within his Lebiitae along with perigonines, lachnophorines, graphipterines, tetragonoderines, cyclosomines (as Masoreini), pentagonicines, and lebiines.

The tribe is represented in North America by six species belonging to the New World genus Colliuris DeGeer, which currently includes about 80 species.

Tribe Calophaenini. This tribe includes two Neotropical genera: Calophaena Klug (48 species) and Calophaenoidea Liebke (one species). Calophaenines have been placed in the tribe Odacanthini (Horn 1881: 147; Bates 1883a: 163; Reichardt 1977: 435; Lorenz 2005: 439), Lachnophorini (Liebherr 1988: 18), or Ctenodactylini (Erwin 1991b: 44; Ball and Bousquet 2000: 107). Liebherr’s (1988) assumption of a relationship with lachnophorines received support from most molecular analyses conducted by Ober and Maddison (2008: 17). The association of calophaenines with ctenodactylines is based on similarity of the adhesive setae under the tarsomeres (Stork in Ball and Bousquet 2000: 107). Until the relationship of calophaenines is better established I prefer to place them in a distinct tribe.

Tribe Ctenodactylini. This tribe is represented only in the New World and includes about 115 species in 18 genera. Most recent authors agree that this group is closely related to the tribe Hexagoniini of the Eastern Hemisphere. Only three species are found north of Mexico and they belong to the genus Leptotrachelus Latreille along with about 30 more species in the tropics.

The main characteristic of the Ctenodactylini and Hexagoniini is the inverted median lobe of the aedeagus as in the drimostomatines.

Tribe Hexagoniini. This tribe contains 65 species in three genera: Hexagonia Kirby (47 species in Asia, Africa, New Guinea [one species], and Australia [one undescribed species, cf. Darlington 1968: 202]), Dinopelma Bates (13 species in the Oriental Region), and Omphreoides Fairmaire (five Madagascan species). Vigna Taglianti and Rossi (1998: 515) indicated that hexagoniines could be related to odacanthines based on similar parasitic laboulbeniales.

Tribe Cyclosomini. This tribe is used here in a restricted sense (see Ball and Bousquet 2000: 109). It includes about 120 species, predominantly tropical, placed in four genera: Mnuphorus Chaudoir with 11 species in the Palaearctic Region; Cyclosomus Latreille with 13 species in the Afrotropical and Oriental Regions; Cyclicus Jeannel with 22 species in the Afrotropical and Oriental Regions; and Tetragonoderus Dejean represented by about 80 species in the Afrotropical, Oriental, Neotropical, and Nearctic Regions, and on the southern fringe of the Palaearctic Region. Several authors, including Jeannel (1949a: 860) and Basilewsky (1984: 527), have considered the New World genus Nemotarsus LeConte as related to cyclosomines, but following Ball (1960b: 157) and Lindroth (1969a: 1014) the genus is listed here in the tribe Lebiini.

Cyclosomines are associated with somoplatines, graphipterines, corsyrines, masoreines, and sarothrocrepidines by most authors based on the presence of long tibial spurs in adults. However, Ball and Bousquet (2000: 109) remarked that the complex as a whole is probably not monophyletic. Molecular analyses published by Ober and Maddison (2008: 17) did not support a close relationships between cyclosomines (sensu lato, i.e., including somoplatines, corsyrines, masoreines, and sarothrocrepidines) and graphipterines but suggested they may be associated with members of dromiine and / or cymindidine lebiines.

Tribe Somoplatini. Ball and Bousquet (2000: 109) restricted this tribe to the genera Somoplatus Dejean (14 Indo-African species), Somoplatodes Basilewsky (two Afrotropical species), and Lophidius Dejean (one Afrotropical species), with Paralophidius Basilewsky recently placed in synonymy with Somoplatus (Schüle 2009: 461). Basilewsky (1986) listed these genera as part of his tribe Masoreini.

Tribe Masoreini. This tribe, as restricted by Ball and Bousquet (2000: 109), comprises the genera Masoreus Dejean (seven Palaearctic species), Atlantomasoreus Mateu (two Moroccan species), Anaulacus Macleay (38 species), and Leuropus Andrewes (one Oriental species). Odontomasoreus Darlington (one species from New Guinea), listed as a distinct genus by Lorenz (2005: 451), has been considered a subgenus of Anaulacus by Ball and Shpeley (2002b: 279). Jeannel (1949a: 860) associated masoreines with cyclosomines and nemotarsines.

Tribe Corsyrini. This tribe comprises the Palaearctic Asian genera Corsyra Dejean (one species) and Discoptera Semenov (five species). Jeannel (1949a: 860) included them with masoreines and Ball and Bousquet (2000: 109) with graphipterines. These authors did not offer evidence to support their groupings.

Tribe Sarothrocrepidini. This tribe is represented by a single genus, Sarothrocrepis Chaudoir, with 26 Indo-Australian species. Jeannel (1949a: 860) associated the genus with graphipterines, masoreines, cyclosomines, and nemotarsines.

Tribe Graphipterini. This tribe, represented in Africa and the Middle East, includes the genera Graphipterus Latreille (about 145 species), Piezia Brullé (18 species), and Trichopiezia Nègre (one species). Jeannel (1949a: 860) associated graphipterines with sarothrocrepidines, masoreines, cyclosomines, and nemotarsines and Kryzhanovskij (1976a: 90) with the same groups with the exception of the nemotarsines. Ober and Maddison (2008: 17) found no support from their molecular analyses for a close relationship between graphipterines and cyclosomines (sensu lato). Instead they found graphipterines to be closely related to pseudomorphines and orthogoniines.

Tribe Lebiini. A markedly complex, worldwide tribe undoubtedly more diverse both in terms of species and lineages in the tropics than in temperate regions. No synapomorphy is known to suggest that this tribe constitutes a monophyletic lineage and Ober and Maddison (2008: 18) did not recover a monophyletic Lebiini in their analyses based on molecular data sequences. The supraspecific classification is not established clearly since most modern studies on lebiines have focussed on regional faunas. The systematic position of some groups within the Lebiini is still debated. For example, the genus Celaenephes Schmidt-Göbel has been considered the most ancestral group of extant lebiines by Shpeley et al. (1985) but as a platynine by Basilewsky (1984). In phylogenetic analyses derived from molecular sequence data by Ober and Maddison (2008: 18), Celaenephes was not clearly associated with members of Lebiini.

Relationships of the tribe are not clearly understood. Jeannel (1948a: 378) included lebiines with anthiines, helluonines, dryptines, galeritines, physocrotaphines, zuphiines, and calophaenines in his superfamily Lebiomorphi. Kryzhanovskij (1976a: 90) used a similar grouping with the exception that he also included orthogoniines, a group that Jeannel (1948a: 377) and Basilewsky (1984: 528) included within the Callistomorphi (i.e., pentagonicines, licinines, panagaeines, chlaeniines, and glyptines). Erwin and Sims (1984: 357) and Erwin (1985: 468) associated lebiines with perigonines, lachnophorines, graphipterines, cyclosomines, masoreines, pentagonicines, and odacanthines.

The 220 genera currently recognized within this tribe are arrayed in the following 16 subtribes (see Ball and Bousquet 2000: 110): Celaenephina, Pericalina (including coptoderines and eucheilines), Sugimotoina, Actenonycina, Apenina, Cymindidina, Dromiusina, Lebiina, Physoderina, Metallicina, Agrina, Calleidina, Gallerucidiina, Peliocypadina, Demetriadina, and Nemotarsina. In the phylogenetic analyses from molecular data published by Ober and Maddison (2008), a small number of clades within the Lebiini were supported but none of these corresponded to the current subtribes, and the subtribes were not recovered as monophyletic.

Tribe Dryptini. Dryptines have been included within the galeritines by several authors (including Darlington 1971: 198). The two lineages are now placed in different tribes but most authors agree that they are closely related. Dryptines and galeritines share some character states with zuphiines and these three groups, referred to as supertribe Zuphiitae by Erwin and Sims (1984: 356) and Erwin (1985: 468), probably constitute a clade. Basilewsky (1960) recognized six dryptine genera in the world, only one of them, the monobasic Amazonian Neodrypta Basilewsky, is found in the New World. Most species are tropically-adapted in the Afrotropical, Australian, and Oriental Regions, with a few species in the southern parts of the Palaearctic Region.

Tribe Galeritini. This tribe is represented in all zoogeographical regions but is more diverse in the tropics than in the temperate zones. Basilewsky (1963b: 7) and Ball (1985) have recognized two lineages within galeritines, treated as subtribes by Ball (1985): Planetina for the genus Planetes Macleay (27 species in the Eastern Hemisphere) and Galeritina for the remaining genera. Lorenz (2005: 507), however, included planetines in zuphiines and Ober and Maddison (2008: Fig.5) found support for such a grouping in some of their analyses and reported that the tribe Galeritini was not monophyletic, based on their molecular data analyses. As indicated previously, this tribe is probably closely related to the Dryptini and Zuphiini.

Tribe Zuphiini. This group is represented in all zoogeographical regions of the world but is more diverse in the tropics than in temperate regions. Zuphiines are grouped by some authors (e.g., Basilewsky 1962a: 100-101) into three subtribes: Leleupidiina, Patriziina, and Zuphiina. Baehr (1985) briefly discussed the status of each of these subtribes. On the other hand, Lorenz (2005: 505-507) included patriziines within the subtribe Zuphiina. Most authors agree that Zuphiini are closely related to Galeritini. For example, LeConte and Horn (1883: 41) and Erwin (1991a: 10) combined zuphiines with galeritines and dryptines. Habu (1967) included zuphiines and galeritines in a single tribe and recognized three subtribes: Zuphiina, Galeritina, and Planetina. Moore (1998: 369) suggested a close relationship between zuphiines and physocrotaphines based on characters of adults and Jeannel (1949a: 1047) enlisted zuphiines, galeritines, dryptines, and physocrotaphines in his family Dryptidae. Ober and Maddison (2008: 18) found a well-supported “Zuphiitae” clade including zuphiines, anthiines, dryptines, galeritines, helluonines, and physocrotaphines in their phylogenetic analyses. However, the tribe Zuphiini was not found to be monophyletic.

The hypogean and monospecific genus Ildobates Español from the Iberian Mountain range was originally described in the Dryptini and was subsequently transferred to the Galeritini by Jeanne (1972) and finally to the Zuphiini by Ortuño et al. (2005).

Tribe Physocrotaphini. The Physocrotaphini includes the following genera: Helluodes Westwood (three species in southwestern India and Sri Lanka), Physocrotaphus Parry (one species from Sri Lanka), Pogonoglossus Chaudoir (35 species), and Schuelea Baehr (three species in New Guinea). The monobasic genus Holoponerus Fairmaire from New Britain was originally described as a lebiine but Darlington (1968) and Moore (1998: 370) agreed that the genus probably belongs to the physocrotaphines. Unfortunately the sole known specimen of Holoponerus godeffroyi Fairmaire was destroyed in 1943 during the bombing of Hamburg in World War II (Moore 1998: 370). All but two species of physocrotaphines are very rarely collected and little is known about their way of life (Sabu et al. 2008: 30). Members of Helluodes are probably termitophilous and those of Pogonoglossus are predominantly subcorticolous though some species could be litter-dwelling forms (Sabu et al. 2008: 41-42).

Jeannel (1949a: 1047) associated physocrotaphines with zuphiines, galeritines, and dryptines and most authors currently agree that these lineages are probably closely related. Jeannel (1949a: 1047) claimed that the genus Pogonoglossus belongs to the Zuphiini near the genus Eunostus Laporte but both Darlington (1968) and Moore (1998: 375) retained the genus within the Physocrotaphini. In his work on the French fauna, Jeannel (1942: 1017) associated physocrotaphines with anthiines and helluonines.

Tribe Anthiini. Anthiines are large, apterous beetles which live in the steppes and subdesert areas of Africa and southwestern Asia. The species are classified into eight or nine genera. The group is clearly defined and probably closely related to the tribe Helluonini (van Emden 1937; Jeannel 1949a: 1040; Bousquet 1987c: 928; Arndt 1993: 44). Basilewsky (1962a: 93) even suggested that anthiines could be derived from a helluonine genus close to Triaenogenius Chaudoir of the Afrotropical Region. Based on larval character states, Bousquet (1987c: 928) suggested that pseudomorphines and/or galeritines could be closely related to the anthiine-helluonine lineage. Erwin and Sims (1984: 356) and Erwin (1985: 468) combined the anthiines with the helluonines and physocrotaphines in their supertribe Anthiitae.

Tribe Helluonini. This is a moderately diverse group with representatives in all continents except Europe and Antarctica. Despite the fact that the group appears homogeneous from the morphology of the adults, Ober and Maddison (2008: 18) did not recover it as monophyletic in their analyses derived from molecular data sequences. Several larval features suggest that helluonines are closely related to anthiines (Bousquet 1987c; Arndt 1998: 186). However, Reichardt (1974, 1977) suggested, based on the structures of the adult mouthparts, that helluonines could be closely related to eucheilines (currently placed within the Lebiini). Liebherr and Will (1998: 145) concluded that Helluonini and Galeritini might be sister-groups based on their possession of a secondary spermathecal gland.

Sloane (1914) and Reichardt (1974) recognized two subtribes among helluonines: Helluonina with representatives in Australia and New Guinea and Helluomorphina (= Omphrina) with representatives in Asia, Africa (including Madagascar), Australia, and the Western Hemisphere. About 165 species, placed in 25 genera, are known worldwide but only eight, all belonging to the New World genus Helluomorphoides Ball, are found in North America.

Tribe Xenaroswellianini. Erwin (2007b: 567) suggested that this recently described tribe, which includes a single species known only from the holotype collected in the Brazilian state of Goiás, could have “a possible relationship with the enigmatic Pseudomorphini.”

Tribe Pseudomorphini. Members of Pseudomorphini are structurally aberrant possibly in response to the group’s evolution into myrmecophily. Nevertheless placement of the group within the subfamily Harpalinae has been confirmed in almost all recent analyses (Arndt et al. 2005: 141). Relationships of pseudomorphines to other harpaline tribes are unsettled as stated by Deuve (1993: 98). Erwin (1981a: 66) remarked that the male genitalia, tarsi, and adult chemical defences suggest that pseudomorphines could be “related somehow to a basal Pterostichitae stock” and that the “paramere vestiture also suggests a connection with Psydritae.” Liebherr and Will (1998: 144) indicated that pseudomorphines, geobaenines, lachnophorines, and odacanthines (including pentagonicines) may be closely related based on the presence of an elongate sclerite joining the spermathecal duct to the common oviduct. Erwin and Stork (1985: 445) concluded that {pseudomorphines + cnemacanthines (= cnemalobines) + scaritines} form a clade that could be the sister-group to Hiletini. Jeannel (1942: 1102; 1949a: 1079) associated pseudomorphines with brachinines under the name Balteifera but almost all authors now agree that the two lineages are not closely related. Ober and Maddison (2008: 18-19) reported strong molecular support for a clade including pseudomorphines, orthogoniines, and graphipterines. They indicated that while morphology does not support such a close relationship, all or some of the members of each tribe have obligate relationships with social insects.

Brief faunistic assessment

The North American fauna currently consists of 2, 676 valid species-group taxa (2, 439 species) of Geadephaga. Of this number, 64 are adventive on this continent, leaving 2, 612 (97.6%) native species-group taxa (2, 375 species).

Adventive species. Table 4 lists the adventive species found on this continent. All but two are accidental introductions (i.e., immigrants). Several carabids were intentionally introduced in New England during the first half of the XX Century for gypsy moth (Lymantria dispar Linnaeus) control, including Calosoma chinense Kirby, Calosoma inquisitor Linnaeus, Calosoma reticulatum Fabricius, Carabus arvensis Herbst, Carabus violaceus Linnaeus, Carabus glabratus Paykull, and Carabus coriaceus Linnaeus (see Smith 1959), but only Calosoma sycophanta and Carabus auratus auratus have become established.

Among the 62 species accidentally introduced, Laemostenus complanatus, Laemostenus terricola terricola, Somotrichus unifasciatus, Plochionus pallens, and Perigona nigriceps are commonly found in cellars or associated with stored products and are now considered subcosmopolitan (Hinton 1945: 21, 27-34). These species have probably been introduced in food and household goods in North America and many other places in the world.

Most of the remaining 57 adventive species found on this continent prior to the XX Century have probably been introduced with bulk rock, bricks, mortar and no doubt soil taken aboard sailing vessels as ballast and dump ashore on this continent (Brown 1940a; Lindroth 1957c). In the XX Century, species have been introduced into North America likely through shipments of nursery stock, within the soil bagged around the roots, as suggested by Kavanaugh and Erwin (1985), Spence and Spence (1988: 166), and Bousquet (1992a). Despite that quarantine regulations were established in the 1960s for the importation of nursery stock, additional species still continue to land and become established on this continent (Fig. 1). The most recent one is Nebria brevicollis in the Willamette Valley in northwestern Oregon (Kavanaugh and LaBonte 2008).

All but four of the adventive species, other than those introduced with stored food products, occur naturally in Europe and their North American populations likely originated from that continent and particularly from southwestern England (Lindroth 1957c: 172). Clivina vespertina and Tetragonoderus laevigatus are native to South America (Nichols 1985b: 380; Shpeley and Ball 2008) while Metacolpodes buchanani (see Habu 1978: 125) and Mochtherus tetraspilotus (see Habu 1982: 87) are native to Asia.

As far as known, none of the adventive species of carabids found in North America are invasive in that they threaten ecosystems, habitats, or even native species.

Figure 1 illustrates the arrival of adventive species in North America through time based on the collecting date of the first inventoried specimen. The rate of arrival has been steady in the XX Century.

Table 4.

List of species-group taxa adventive and established in North America.

Species Introduced in YFC
East West South
Leistus ferrugineus (Linnaeus)     1977
Nebria brevicollis (Fabricius)     2007
Notiophilus biguttatus (Fabricius)   1923
Notiophilus palustris (Duftschmid)     1968
Calosoma sycophanta (Linnaeus)*     N/A
Carabus granulatus granulatus Linnaeus   1890
Carabus nemoralis nemoralis O.F. Müller   1890
Carabus auratus auratus Linnaeus*     N/A
Clivina vespertina Putzeys     1948
Clivina collaris (Herbst)   <1838
Clivina fossor fossor (Linnaeus)   1915
Dyschirius globosus Herbst     1978
Broscus cephalotes (Linnaeus)     1987
Blemus discus discus (Fabricius)     1933
Trechus obtusus Erichson     1927
Trechus quadristriatus (Schrank)     1965
Trechus rubens (Fabricius)     <1863
Asaphidion curtum curtum (Heyden)     1930
Bembidion lampros (Herbst)   1947
Bembidion properans (Stephens)     1942
Bembidion obtusum Audinet-Serville     1956
Bembidion nigropiceum (Marsham)     <1897
Bembidion stephensii Crotch     1891
Bembidion bruxellense Wesmael     1907
Bembidion femoratum femoratum Sturm     1967
Bembidion tetracolum tetracolum Say   <1823
Elaphropus parvulus (Dejean)     1940
Porotachys bisulcatus (Nicolai)   <1900
Stomis pumicatus (Panzer)     1984
Pterostichus vernalis (Panzer)     1997
Pterostichus strenuus (Panzer)   1937
Pterostichus melanarius melanarius (Illiger)   1926
Abax parallelepipedus (Piller & Mitterpacher)     1965
Amara aulica (Panzer)     1929
Amara apricaria (Paykull) ?   <1865
Amara fulva (O.F. Müller)     1905
Amara bifrons (Gyllenhal)     1929
Amara ovata (Fabricius)   1925
Amara aenea (DeGeer)   1904
Amara anthobia Villa & Villa   1929
Amara communis (Panzer)     1988
Amara eurynota (Panzer)     1971
Amara familiaris (Duftschmid)   1901
Anisodactylus binotatus (Fabricius)     1911
Bradycellus harpalinus (Audinet-Serville)     1951
Acupalpus meridianus (Linnaeus)   1931
Ophonus puncticeps Stephens     1954
Ophonus rufibarbis (Fabricius)     1953
Harpalus rufipes (DeGeer)     1937
Harpalus affinis (Schrank)   <1798
Harpalus rubripes (Duftschmid)     1981
Calathus fuscipes (Goeze)     1928
Laemostenus complanatus (Dejean)     <1874
Laemostenus terricola terricola (Herbst) ?   <1894
Paranchus albipes (Fabricius)     <1835
Agonum muelleri (Herbst)   1840
Metacolpodes buchanani (Hope)     1931
Perigona nigriceps (Dejean)   <1853
Tetragonoderus laevigatus Chaudoir     2007
Mochtherus tetraspilotus (Macleay)     1992
Somotrichus unifasciatus (Dejean)     ?
Dromius fenestratus (Fabricius)     1952
Philorhizus melanocephalus (Dejean)     1996
Plochionus pallens (Fabricius) ? <1833

YFC: Year of first collection

Native species. The known North American fauna contains 2, 612 native species-group taxa (2, 375 species), of which 2, 071 are endemic (79.3%). Among the 541 non-endemic species-group taxa, 98 (3.7% of the entire native fauna) are Holarctic and 446 (17.1%) extend south of North America as defined in this catalogue.

The number of 2, 612 native species-group taxa obviously does not give an accurate account of the size of the North American carabid fauna. Several factors are involved. First, 65 species-group taxa described by Casey have not been treated subsequently. Considering that only 13.1% of Casey’s North American species-group taxa subsequently treated are currently valid, I estimate that about 50 of the remaining Casey taxa will eventually be synonymized. This would bring the number of valid Nearctic species-group taxa to about 2, 560 (2, 325 species). Second, several genus-group taxa have not been revised in “modern times” and obviously the number of valid species-group taxa will change. The following generic taxa are those that need revisions: Clivina Latreille (except Antroforceps), Paraclivina Kult, Elaphropus Motschulsky, Tachys Dejean, Poecilus Bonelli, Hypherpes Chaudoir, Stenocellus Casey, Selenophorus Dejean, Discoderus LeConte, Olisthopus Dejean, Pinacodera Schaum, Apristus Chaudoir, Axinopalpus LeConte, Zuphium Latreille, Pseudaptinus Laporte, and Pseudomorpha Kirby. Third, many species-group taxa are known but still undescribed. For example, Barr (2004: 1) reported that he was aware of 80 undescribed species of Pseudanophthalmus. Fourth, through modern techniques, such as DNA sequences, several “species” will probably be found to be a complex of two or more species. For example, Maddison (2008) found out that Lindroth’s (1963b) concept of Bembidion chalceum and Bembidion honestum was a complex of seven species. Finally, there is no doubt that many species remain to be discovered in North America, especially in peculiar habitats, such as caves, burrows, and deep litter.

Figure 2 illustrates through decades the number of North American species-group taxa described and the number of those regarded as valid today based on the present catalogue. It shows that the periods between 1821-1880, 1911-1930, and 1961-1990 have been the most prolific. The first one, 1821-1880, can be associated with the works of John L. LeConte (724 geadephagan species-group taxa described), Count Dejean (289 taxa), Thomas Say (164 taxa), Baron de Chaudoir (126 taxa), and Victor Motschulsky (121 taxa). The second period, 1911-1930, relates of course to the work of Colonel Casey. By the time of his last publication in 1924, Casey had described 1, 864 species-group taxa of carabids based on North American specimens while the number of taxa proposed by all authors combined since Linnaeus amounted to 2, 288. The third one, 1961-1990, is connected to the magnificent work of Carl H. Lindroth (76 taxa) on the Carabidae of Canada and Alaska, the work of Thomas Barr (147 taxa) on the cave fauna, and the leadership of George Ball who, besides his own research, directed many students.

Figure 3 shows the growth, through time, of the number of species-group taxa described from North American specimens and those still considered valid. Taking into consideration the trend of description of species-group taxa still valid, the number of native species-group taxa should be slightly over 3, 000 by the year 2060.

Figure 1.

Number of adventive species of Geadephaga established in North America through time.

Figure 2.

Number of North American species-group taxa of Geadephaga described and currently valid, by decade.

Figure 3.

Cumulative number of species-group taxa of Geadephaga described and currently valid, by decade.

Political regions. Table 5 gives statistics regarding the number of geadephagan species-group taxa for each political region covered in this catalogue.

Texas (693 native species-group taxa), California (646 taxa), North Carolina (536 taxa), New York (528 taxa), and Virginia (520 taxa) have the highest number of native taxa recorded and Greenland (4 native taxa), Nunavut (34 taxa), Saint Pierre and Miquelon (50 taxa), Labrador (91 taxa), and Prince Edward Island (144 taxa) the lowest. Are these numbers representative of the relative sizes of the regions? Only to some extent. Some regions, for example Nunavut and Labrador, have been much less surveyed than others like Newfoundland proper. The geadephagan fauna of most states are still inadequately known. In the past 50 years, only Maine (Majka et al. 2011; Bousquet 2010a), New Hampshire and Vermont (Bousquet 2010a), Wisconsin (Messer 2010), Rhode Island (Sikes 2004), Connecticut (Krinsky and Oliver 2001), South Carolina (Ciegler 2000), Florida (Peck and Thomas 1998), South Dakota (Kirk and Balsbaugh 1975), and Alaska (Lindroth 1961-1969) have been the subject of faunistic assessments (see Table 3). Several states never had a checklist of their geadephagan fauna published.

Table 5.

Species-group taxon statistics for political regions.

Political regions NbNT NbAd %NF Rank
Alabama 439 4 16.8 22
Alaska 248 2 09.5 56
Alberta 405 11 15.5 32
Arizona 502 3 19.2 9
Arkansas 403 4 15.5 33
British Columbia 479 28 18.3 12
California 646 12 24.7 2
Colorado 450 3 17.2 20
Connecticut 383 23 14.7 39
Delaware 201 5 07.7 59
District of Columbia 337 6 12.9 48
Florida 383 8 14.7 39
Georgia 478 3 18.3 14
Greenland 4 0 0.2 66
Idaho 338 10 12.9 47
Illinois 504 9 19.3 8
Indiana 479 9 18.3 12
Iowa 431 9 16.5 25
Kansas 408 4 15.6 30
Kentucky 371 2 14.2 42
Labrador 91 4 03.5 63
Louisiana 399 4 15.3 34
Maine 383 27 14.7 39
Manitoba 370 6 14.1 43
Maryland 408 9 15.5 30
Massachusetts 426 27 16.3 27
Michigan 474 15 18.2 16
Minnesota 419 10 16.0 28
Mississippi 439 5 16.8 22
Missouri 396 5 15.1 36
Montana 358 6 13.6 45
Nebraska 358 4 13.7 45
Nevada 239 3 09.2 57
New Brunswick 302 30 11.6 52
Newfoundland 151 27 05.8 61
New Hampshire 429 25 16.4 26
New Jersey 456 10 17.5 18
New Mexico 454 0 17.3 19
New York 528 21 20.2 4
North Carolina 536 4 20.4 3
North Dakota 275 4 10.5 54
Northwest Territories 218 1 08.3 58
Nova Scotia 258 35 09.8 55
Nunavut 34 0 01.3 65
Ohio 499 16 19.1 11
Oklahoma 387 2 14.8 37
Ontario 510 21 19.5 6
Oregon 478 22 18.2 14
Pennsylvania 500 19 19.1 10
Prince Edward Island 144 29 05.5 62
Quebec 446 32 17.1 21
Rhode Island 300 17 11.5 53
Saint Pierre and Miquelon 50 15 01.9 64
Saskatchewan 336 6 12.9 49
South Carolina 414 4 15.8 29
South Dakota 384 4 14.7 38
Tennessee 506 2 19.3 7
Texas 693 1 26.5 1
Utah 323 5 12.4 50
Vermont 397 21 15.2 35
Virginia 520 11 19.9 5
Washington 440 24 16.8 22
West Virginia 367 7 14.1 44
Wisconsin 463 16 17.7 17
Wyoming 309 4 11.8 51
Yukon Territory 201 0 07.7 59

NbNT = Number of native species-group taxa

NbAd = Number of adventive species-group taxa

%NF = % of the native North American fauna

Table 6.

Checklists and taxonomic treatments on Geadephaga for North American political regions.

Political regions References
Alabama Löding (1945)
Alaska Hamilton (1894a); Lindroth (1961-1969); Bousquet (1991b)
Alberta Lindroth (1961-1969); Bousquet (1991b)
British Columbia Hatch (1953); Lindroth (1961-1969); Bousquet (1991b)
Colorado Wickham (1902)
Connecticut Britton (1920); Krinsky and Oliver (2001)
District of Columbia Ulke (1902)
Florida Schwarz (1878); Leng (1915); Choate (1990); Peck and Thomas (1998)
Georgia Fattig (1949)
Greenland Henriksen (1939); Böcher (1988)
Idaho Hatch (1953)
Indiana Blatchley (1910)
Iowa Wickham (1911b); Jaques and Redlinger (1946)
Kansas Popenoe (1877)
Labrador Sherman (1910); Lindroth (1961-1969); Bousquet (1991b); Bousquet (2010a)
Maine Bousquet (2010a); Majka et al. (2011)
Manitoba Lindroth (1961-1969); Bousquet (1991b)
Massachusetts Harris (1833); Harris (1835)
New Brunswick Lindroth (1961-1969); Bousquet (1991b); Majka et al. (2007); Bousquet (2010a)
Newfoundland Lindroth (1955a); Lindroth (1961-1969); Bousquet (1991b); Bousquet (2010a)
New Hampshire Bousquet (2010a)
New Jersey Smith (1890); Smith (1900); Smith (1910)
New Mexico Fall and Cockerell (1907)
New York Notman (1928)
North Carolina Brimley (1938)
Northwest Territories Lindroth (1961-1969); Bousquet (1991b)
Nova Scotia Lindroth (1961-1969); Lindroth (1954c); Bousquet (1991b); Majka et al. (2007); Bousquet (2010a)
Ontario Lindroth (1961-1969); Bousquet (1991b)
Oregon Hatch (1953)
Prince Edward Island Lindroth (1961-1969); Bousquet (1991b); Majka et al. (2007); Majka et al. (2008); Bousquet (2010a)
Quebec Chagnon (1917); Lindroth (1961-1969); Larochelle (1975); Larochelle (1976); Larochelle (1979); Bousquet (1991b); Laplante et al. (1991); Bousquet (2010a)
Rhode Island Davis (1904); Sikes (2004)
Saint Pierre and Miquelon Lindroth (1955a); Roux (1984); Bousquet (2010a)
Saskatchewan Lindroth (1961-1969); Bousquet (1991b)
South Carolina Kirk (1969 and 1970); Ciegler (2000)
South Dakota Kirk and Balsbaugh (1975)
Vermont Bousquet (2010a)
Washington Hatch (1939b); Hatch (1953)
Wisconsin Messer (2010)
Yukon Territory Lindroth (1961-1969); Bousquet (1991b); Ball and Currie (1997)
List of North American valid species-group taxa 6 Trachypachini

Trachypachus gibbsii LeConte, 1861

Trachypachus inermis Motschulsky, 1850

Trachypachus slevini Van Dyke, 1925


Clinidium (Arctoclinidium) apertum allegheniense Bell & Bell, 1975

Clinidium (Arctoclinidium) apertum apertum Reitter, 1880

Clinidium (Arctoclinidium) baldufi Bell, 1970

Clinidium (Arctoclinidium) calcaratum LeConte, 1875

Clinidium (Arctoclinidium) rosenbergi Bell, 1970

Clinidium (Arctoclinidium) sculptile (Newman, 1838)

Clinidium (Arctoclinidium) valentinei Bell, 1970


Omoglymmius (Boreoglymmius) americanus (Laporte, 1836)

Omoglymmius (Boreoglymmius) hamatus (LeConte, 1875)


Pelophila borealis (Paykull, 1790)

Pelophila rudis (LeConte, 1863)


Opisthius richardsoni Kirby, 1837


Leistus (Leistus) ferrugineus (Linnaeus, 1758)

Leistus (Neoleistus) ferruginosus Mannerheim, 1843

Leistus (Neoleistus) longipennis Casey, 1920

Leistus (Neoleistus) madmeridianus Erwin, 1970

Nippononebria (Vancouveria) altisierrae (Kavanaugh, 1984)

Nippononebria (Vancouveria) campbelli (Kavanaugh, 1984)

Nippononebria (Vancouveria) virescens (Horn, 1870)

Nebria (Boreonebria) bellorum Kavanaugh, 1979

Nebria (Boreonebria) crassicornis crassicornis Van Dyke, 1925

Nebria (Boreonebria) crassicornis intermedia Van Dyke, 1949

Nebria (Boreonebria) frigida Sahlberg, 1844

Nebria (Boreonebria) gouleti Kavanaugh, 1979

Nebria (Boreonebria) gyllenhali castanipes (Kirby, 1837)

Nebria (Boreonebria) gyllenhali lassenensis Kavanaugh, 1979

Nebria (Boreonebria) gyllenhali lindrothi Kavanaugh, 1979

Nebria (Boreonebria) hudsonica LeConte, 1863

Nebria (Boreonebria) lacustris Casey, 1913

Nebria (Boreonebria) nivalis gaspesiana Kavanaugh, 1979

Nebria (Boreonebria) nivalis nivalis (Paykull, 1790)

Nebria (Nakanebria) paradisi Darlington, 1931

Nebria (Nakanebria) turmaduodecima Kavanaugh, 1981

Nebria (Reductonebria) acuta acuta Lindroth, 1961

Nebria (Reductonebria) acuta quileute Kavanaugh, 1979

Nebria (Reductonebria) acuta sonorae Kavanaugh, 1981

Nebria (Reductonebria) appalachia Darlington, 1932

Nebria (Reductonebria) arkansana arkansana Casey, 1913

Nebria (Reductonebria) arkansana edwardsi Kavanaugh, 1979

Nebria (Reductonebria) arkansana fragilis Casey, 1924

Nebria (Reductonebria) arkansana oowah Kavanaugh, 1979

Nebria (Reductonebria) charlottae Lindroth, 1961

Nebria (Reductonebria) danmanni Kavanaugh, 1981

Nebria (Reductonebria) darlingtoni Kavanaugh, 1979

Nebria (Reductonebria) desolata Kavanaugh, 1971

Nebria (Reductonebria) diversa LeConte, 1863

Nebria (Reductonebria) eschscholtzii Ménétriés, 1843

Nebria (Reductonebria) georgei Kavanaugh, 2008

Nebria (Reductonebria) gregaria Fischer von Waldheim, 1820

Nebria (Reductonebria) haida Kavanaugh, 1984

Nebria (Reductonebria) jeffreyi Kavanaugh, 1984

Nebria (Reductonebria) lituyae Kavanaugh, 1979

Nebria (Reductonebria) louiseae Kavanaugh, 1984

Nebria (Reductonebria) lyelli Van Dyke, 1925

Nebria (Reductonebria) mannerheimii Fischer von Waldheim, 1828

Nebria (Reductonebria) navajo Kavanaugh, 1979

Nebria (Reductonebria) obliqua chuskae Kavanaugh, 1979

Nebria (Reductonebria) obliqua obliqua LeConte, 1867

Nebria (Reductonebria) pallipes Say, 1823

Nebria (Reductonebria) sahlbergii modoc Kavanaugh, 1979

Nebria (Reductonebria) sahlbergii sahlbergii Fischer von Waldheim, 1828

Nebria (Reductonebria) sahlbergii triad Kavanaugh, 1979

Nebria (Reductonebria) suturalis LeConte, 1850

Nebria (Reductonebria) wallowae Kavanaugh, 1984

Nebria (Reductonebria) zioni oasis Kavanaugh, 1979

Nebria (Reductonebria) zioni zioni Van Dyke, 1943

Nebria (Catonebria) calva Kavanaugh, 1984

Nebria (Catonebria) carri Kavanaugh, 1979

Nebria (Catonebria) catenata Casey, 1913

Nebria (Catonebria) coloradensis Van Dyke, 1943

Nebria (Catonebria) gebleri albimontis Kavanaugh, 1984

Nebria (Catonebria) gebleri cascadensis Kavanaugh, 1979

Nebria (Catonebria) gebleri fragariae Kavanaugh, 1979

Nebria (Catonebria) gebleri gebleri Dejean, 1831

Nebria (Catonebria) gebleri rathvoni LeConte, 1853

Nebria (Catonebria) gebleri siskiyouensis Kavanaugh, 1979

Nebria (Catonebria) ingens ingens Horn, 1870

Nebria (Catonebria) ingens riversi Van Dyke, 1925

Nebria (Catonebria) kincaidi balli Kavanaugh, 1979

Nebria (Catonebria) kincaidi kincaidi Schwarz, 1900

Nebria (Catonebria) labontei Kavanaugh, 1984

Nebria (Catonebria) meanyi giulianii Kavanaugh, 1981

Nebria (Catonebria) meanyi lamarckensis Kavanaugh, 1979

Nebria (Catonebria) meanyi meanyi Van Dyke, 1925

Nebria (Catonebria) meanyi sylvatica Kavanaugh, 1979

Nebria (Catonebria) metallica Fischer von Waldheim, 1820

Nebria (Catonebria) ovipennis LeConte, 1878

Nebria (Catonebria) piperi Van Dyke, 1925

Nebria (Catonebria) piute piute Erwin & Ball, 1972

Nebria (Catonebria) piute sevieri Kavanaugh, 1984

Nebria (Catonebria) piute utahensis Kavanaugh, 1979

Nebria (Catonebria) praedicta Kavanaugh & Schoville, 2009

Nebria (Catonebria) purpurata LeConte, 1878

Nebria (Catonebria) schwarzi beverlianna Kavanaugh, 1979

Nebria (Catonebria) schwarzi schwarzi Van Dyke, 1925

Nebria (Catonebria) sierrablancae Kavanaugh, 1984

Nebria (Catonebria) spatulata sierrae Kavanaugh, 1979

Nebria (Catonebria) spatulata spatulata Van Dyke, 1925

Nebria (Catonebria) steensensis Kavanaugh, 1984

Nebria (Catonebria) trifaria pasquineli Kavanaugh, 1984

Nebria (Catonebria) trifaria trifaria LeConte, 1878

Nebria (Catonebria) vandykei vandykei Bänninger, 1928

Nebria (Catonebria) vandykei wyeast Kavanaugh, 1979

Nebria (Nebria) brevicollis (Fabricius, 1792)


Notiophilus aeneus (Herbst, 1806)

Notiophilus aquaticus (Linnaeus, 1758)

Notiophilus biguttatus (Fabricius, 1779)

Notiophilus borealis Harris, 1869

Notiophilus directus Casey, 1920

Notiophilus intermedius Lindroth, 1955

Notiophilus nemoralis Fall, 1906

Notiophilus nitens LeConte, 1857

Notiophilus novemstriatus LeConte, 1847

Notiophilus palustris (Duftschmid, 1812)

Notiophilus semiopacus Eschscholtz, 1833

Notiophilus semistriatus Say, 1823

Notiophilus sierranus Casey, 1920

Notiophilus simulator Fall, 1906

Notiophilus sylvaticus Dejean, 1831


Sphaeroderus bicarinatus (LeConte, 1853)

Sphaeroderus canadensis canadensis Chaudoir, 1861

Sphaeroderus canadensis lengi Darlington, 1933

Sphaeroderus indianae (Blatchley, 1910)

Sphaeroderus nitidicollis Guérin-Méneville, 1829

Sphaeroderus schaumii Chaudoir, 1861

Sphaeroderus stenostomus lecontei Dejean, 1826

Sphaeroderus stenostomus stenostomus (Weber, 1801)

Cychrus hemphillii hemphillii Horn, 1879

Cychrus hemphillii rickseckeri LeConte, 1884

Cychrus tuberculatus Harris, 1839

Scaphinotus (Scaphinotus) elevatus coloradensis Van Dyke, 1907

Scaphinotus (Scaphinotus) elevatus elevatus (Fabricius, 1787)

Scaphinotus (Scaphinotus) elevatus flammeus Haldeman, 1844

Scaphinotus (Scaphinotus) elevatus lengi Van Dyke, 1938

Scaphinotus (Scaphinotus) elevatus neomexicanus Van Dyke, 1924

Scaphinotus (Scaphinotus) elevatus tenebricosus Roeschke, 1907

Scaphinotus (Scaphinotus) kelloggi (Dury, 1912)

Scaphinotus (Scaphinotus) parisiana Allen & Carlton, 1988

Scaphinotus (Scaphinotus) petersi biedermani Roeschke, 1907

Scaphinotus (Scaphinotus) petersi catalinae Van Dyke, 1924

Scaphinotus (Scaphinotus) petersi corvus (Fall, 1910)

Scaphinotus (Scaphinotus) petersi grahami Van Dyke, 1938

Scaphinotus (Scaphinotus) petersi kathleenae Ball, 1966

Scaphinotus (Scaphinotus) petersi petersi Roeschke, 1907

Scaphinotus (Scaphinotus) snowi roeschkei Van Dyke, 1907

Scaphinotus (Scaphinotus) snowi snowi (LeConte, 1881)

Scaphinotus (Scaphinotus) unicolor (Fabricius, 1787)

Scaphinotus (Scaphinotus) vandykei Roeschke, 1907

Scaphinotus (Irichroa) irregularis (Beutenmüller, 1903)

Scaphinotus (Irichroa) viduus (Dejean, 1826)

Scaphinotus (Irichroa) webbi Bell, 1959

Scaphinotus (Nomaretus) bilobus (Say, 1823)

Scaphinotus (Nomaretus) cavicollis (LeConte, 1859)

Scaphinotus (Nomaretus) fissicollis (LeConte, 1853)

Scaphinotus (Nomaretus) infletus Allen & Carlton, 1988

Scaphinotus (Nomaretus) liebecki Van Dyke, 1936

Scaphinotus (Maronetus) debilis alpinus (Beutenmüller, 1903)

Scaphinotus (Maronetus) debilis debilis (LeConte, 1853)

Scaphinotus (Maronetus) hoffmani (Barr, 2009)

Scaphinotus (Maronetus) hubbardi (Schwarz, 1895)

Scaphinotus (Maronetus) imperfectus (Horn, 1861)

Scaphinotus (Maronetus) incompletus (Schwarz, 1895)

Scaphinotus (Maronetus) reichlei (Barr, 2009)

Scaphinotus (Maronetus) schwarzi (Beutenmüller, 1913)

Scaphinotus (Maronetus) tenuis (Casey, 1914)

Scaphinotus (Maronetus) unistriatus (Darlington, 1932)

Scaphinotus (Steniridia) aeneicollis (Beutenmüller, 1903)

Scaphinotus (Steniridia) andrewsii amplicollis (Casey, 1920)

Scaphinotus (Steniridia) andrewsii andrewsii (Harris, 1839)

Scaphinotus (Steniridia) andrewsii darlingtoni (Valentine, 1935)

Scaphinotus (Steniridia) andrewsii germari (Chaudoir, 1861)

Scaphinotus (Steniridia) andrewsii mutabilis (Casey, 1920)

Scaphinotus (Steniridia) andrewsii parvitarsalis (Valentine, 1935)

Scaphinotus (Steniridia) andrewsii waldensius (Valentine, 1935)

Scaphinotus (Steniridia) guyotii (LeConte, 1863)

Scaphinotus (Steniridia) lodingi lodingi (Valentine, 1935)

Scaphinotus (Steniridia) lodingi obscurus (Valentine, 1935)

Scaphinotus (Steniridia) ridingsii monongahelae Leng, 1917

Scaphinotus (Steniridia) ridingsii ridingsii (Bland, 1863)

Scaphinotus (Steniridia) tricarinatus (Casey, 1914)

Scaphinotus (Steniridia) violaceus (LeConte, 1863)

Scaphinotus (Pseudonomaretus) mannii Wickham, 1919

Scaphinotus (Pseudonomaretus) merkelii (Horn, 1890)

Scaphinotus (Pseudonomaretus) regularis (LeConte, 1884)

Scaphinotus (Pseudonomaretus) relictus (Horn, 1881)

Scaphinotus (Stenocantharus) angusticollis (Mannerheim, 1823)

Scaphinotus (Stenocantharus) hatchi Beer, 1971

Scaphinotus (Stenocantharus) johnsoni Van Dyke, 1924

Scaphinotus (Stenocantharus) velutinus (Ménétriés, 1843)

Scaphinotus (Brennus) bullatus Van Dyke, 1924

Scaphinotus (Brennus) cordatus (LeConte, 1853)

Scaphinotus (Brennus) crenatus (Motschulsky, 1859)

Scaphinotus (Brennus) cristatus (Harris, 1839)

Scaphinotus (Brennus) interruptus (Ménétriés, 1843)

Scaphinotus (Brennus) marginatus (Fischer von Waldheim, 1820)

Scaphinotus (Brennus) obliquus (LeConte, 1868)

Scaphinotus (Brennus) oreophilus (Rivers, 1890)

Scaphinotus (Brennus) punctatus (LeConte, 1859)

Scaphinotus (Brennus) riversi (Roeschke, 1907)

Scaphinotus (Brennus) rugiceps incipiens (Casey, 1897)

Scaphinotus (Brennus) rugiceps rugiceps (Horn, 1872)

Scaphinotus (Brennus) striatopunctatus (Chaudoir, 1844)

Scaphinotus (Brennus) subtilis (Schaum, 1863)

Scaphinotus (Brennus) ventricosus (Dejean, 1831)

Scaphinotus (Neocychrus) angulatus (Harris, 1839)

Scaphinotus (Neocychrus) behrensi (Roeschke, 1907)

Scaphinotus (Neocychrus) longiceps Van Dyke, 1924


Calosoma (Castrida) sayi Dejean, 1826

Calosoma (Calosoma) frigidum Kirby, 1837

Calosoma (Calosoma) sycophanta (Linnaeus, 1758)

Calosoma (Calodrepa) aurocinctum Chaudoir, 1850

Calosoma (Calodrepa) scrutator (Fabricius, 1775)

Calosoma (Calodrepa) splendidum Dejean, 1831

Calosoma (Calodrepa) wilcoxi LeConte, 1847

Calosoma (Camegonia) marginale Casey, 1897

Calosoma (Camegonia) parvicolle Fall, 1910

Calosoma (Camegonia) prominens LeConte, 1853

Calosoma (Carabosoma) angulatum Chevrolat, 1834

Calosoma (Carabosoma) eremicola Fall, 1910

Calosoma (Carabosoma) peregrinator Guérin-Méneville, 1844

Calosoma (Carabosoma) sponsa Casey, 1897

Calosoma (Callitropa) externum (Say, 1823)

Calosoma (Callitropa) macrum LeConte, 1853

Calosoma (Callitropa) protractum LeConte, 1862

Calosoma (Blaptosoma) haydeni haydeni Horn, 1870

Calosoma (Blaptosoma) haydeni punctulicolle Bates, 1891

Calosoma (Chrysostigma) affine Chaudoir, 1843

Calosoma (Chrysostigma) calidum (Fabricius, 1775)

Calosoma (Chrysostigma) cancellatum Eschscholtz, 1833

Calosoma (Chrysostigma) lepidum LeConte, 1845

Calosoma (Chrysostigma) morrisonii Horn, 1885

Calosoma (Chrysostigma) obsoletum Say, 1823

Calosoma (Chrysostigma) semilaeve LeConte, 1852

Calosoma (Chrysostigma) simplex LeConte, 1878

Calosoma (Chrysostigma) tepidum LeConte, 1852

Calosoma (Callistenia) dawsoni (Dajoz, 1997)

Calosoma (Callistenia) dietzii Schaeffer, 1904

Calosoma (Callistenia) discors LeConte, 1857

Calosoma (Callistenia) lariversi Van Dyke, 1943

Calosoma (Callistenia) latipenne Horn, 1870

Calosoma (Callistenia) luxatum Say, 1823

Calosoma (Callistenia) moniliatum (LeConte, 1852)

Calosoma (Callistenia) monticola Casey, 1897

Calosoma (Callistenia) oregonum (Gidaspow, 1959)

Calosoma (Callistenia) placerum (Gidaspow, 1959)

Calosoma (Callistenia) schaefferi Breuning, 1928

Calosoma (Callistenia) subaeneum Chaudoir, 1869

Calosoma (Callistenia) subasperatum Schaeffer, 1915

Calosoma (Callistenia) wilkesii (LeConte, 1852)

Carabus (Carabus) goryi Dejean, 1831

Carabus (Carabus) granulatus granulatus Linnaeus, 1758

Carabus (Carabus) vinctus (Weber, 1801)

Carabus (Diocarabus) chamissonis Fischer von Waldheim, 1820

Carabus (Aulonocarabus) truncaticollis truncaticollis Eschscholtz, 1833

Carabus (Homoeocarabus) maeander maeander Fischer von Waldheim, 1820

Carabus (Hemicarabus) serratus Say, 1823

Carabus (Archicarabus) nemoralis nemoralis Müller, 1764

Carabus (Tachypus) auratus auratus Linnaeus, 1760

Carabus (Tanaocarabus) finitimus Haldeman, 1852

Carabus (Tanaocarabus) forreri forreri Bates, 1882

Carabus (Tanaocarabus) sylvosus Say, 1823

Carabus (Tanaocarabus) taedatus agassii LeConte, 1850

Carabus (Tanaocarabus) taedatus bicanaliceps Casey, 1920

Carabus (Tanaocarabus) taedatus rainieri Van Dyke, 1945

Carabus (Tanaocarabus) taedatus taedatus Fabricius, 1787

Carabus (Megodontus) vietinghoffii vietinghoffii Adams, 1812


Amblycheila baroni Rivers, 1890

Amblycheila cylindriformis (Say, 1823)

Amblycheila hoversoni Gage, 1991

Amblycheila picolominii Reiche, 1840

Amblycheila schwarzi Horn, 1904

Omus audouini Reiche, 1838

Omus californicus angustocylindricus Horn, 1913

Omus californicus californicus Eschscholtz, 1829

Omus californicus intermedius Leng, 1902

Omus californicus subcylindricus Nunenmacher, 1940

Omus cazieri van den Berghe, 1994

Omus dejeanii Reiche, 1838

Omus submetallicus Horn, 1869


Tetracha (Tetracha) carolina carolina (Linnaeus, 1763)

Tetracha (Tetracha) floridana Leng & Mutchler, 1916

Tetracha (Tetracha) virginica (Linnaeus, 1767)

Tetracha (Neotetracha) impressa (Chevrolat, 1841)


Cylindera (Cylindera) celeripes (LeConte, 1846)

Cylindera (Cylindera) cursitans (LeConte, 1856)

Cylindera (Cylindera) debilis (Bates, 1890)

Cylindera (Cylindera) lemniscata lemniscata (LeConte, 1854)

Cylindera (Cylindera) lemniscata rebaptisata (Vaurie, 1951)

Cylindera (Cylindera) lunalonga (Schaupp, 1884)

Cylindera (Cylindera) terricola cinctipennis (LeConte, 1846)

Cylindera (Cylindera) terricola continua (Knaus, 1923)

Cylindera (Cylindera) terricola imperfecta (LeConte, 1851)

Cylindera (Cylindera) terricola kaibabensis (Johnson, 1990)

Cylindera (Cylindera) terricola susanagreae (Kippenhan, 2007)

Cylindera (Cylindera) terricola terricola (Say, 1824)

Cylindera (Cylindera) unipunctata (Fabricius, 1775)

Cylindera (Cylindera) viridisticta arizonensis (Bates, 1884)

Ellipsoptera blanda (Dejean, 1831)

Ellipsoptera cuprascens (LeConte, 1852)

Ellipsoptera gratiosa (Guérin-Méneville, 1840)

Ellipsoptera hamata lacerata (Chaudoir, 1854)

Ellipsoptera hamata monti (Vaurie, 1951)

Ellipsoptera hirtilabris (LeConte, 1875)

Ellipsoptera lepida (Dejean, 1831)

Ellipsoptera macra ampliata (Vaurie, 1951)

Ellipsoptera macra fluviatilis (Vaurie, 1951)

Ellipsoptera macra macra (LeConte, 1856)

Ellipsoptera marginata (Fabricius, 1775)

Ellipsoptera nevadica citata (Rumpp, 1977)

Ellipsoptera nevadica knausii (Leng, 1902)

Ellipsoptera nevadica lincolniana (Casey, 1916)

Ellipsoptera nevadica makosika (Spomer, 2004)

Ellipsoptera nevadica nevadica (LeConte, 1875)

Ellipsoptera nevadica olmosa (Vaurie, 1951)

Ellipsoptera nevadica tubensis (Cazier, 1939)

Ellipsoptera puritana (Horn, 1871)

Ellipsoptera rubicunda (Harris, 1911)

Ellipsoptera sperata inquisitor (Casey, 1897)

Ellipsoptera sperata sperata (LeConte, 1856)

Ellipsoptera wapleri (LeConte, 1875)

Microthylax olivaceus (Chaudoir, 1854)

Opilidia chlorocephala smythi (Harris, 1913)

Brasiella wickhami (Horn, 1903)

Dromochorus belfragei Sallé, 1877

Dromochorus pilatei Guérin-Méneville, 1849

Dromochorus pruininus Casey, 1897

Dromochorus velutinigrens Johnson, 1992

Habroscelimorpha californica mojavi (Cazier, 1937)

Habroscelimorpha californica pseudoerronea (Rumpp, 1958)

Habroscelimorpha circumpicta circumpicta (LaFerté-Sénectère, 1841)

Habroscelimorpha circumpicta johnsonii (Fitch, 1857)

Habroscelimorpha circumpicta pembina (Johnson, 1993)

Habroscelimorpha dorsalis dorsalis (Say, 1817)

Habroscelimorpha dorsalis media (LeConte, 1856)

Habroscelimorpha dorsalis saulcyi (Guérin-Méneville, 1840)

Habroscelimorpha dorsalis venusta (LaFerté-Sénectère, 1841)

Habroscelimorpha fulgoris albilata (Acciavatti, 1981)

Habroscelimorpha fulgoris erronea (Vaurie, 1951)

Habroscelimorpha fulgoris fulgoris (Casey, 1913)

Habroscelimorpha gabbii (Horn, 1867)

Habroscelimorpha pamphila (LeConte, 1873)

Habroscelimorpha praetextata pallidofemora (Acciavatti, 1981)

Habroscelimorpha praetextata praetextata (LeConte, 1854)

Habroscelimorpha severa (LaFerté-Sénectère, 1841)

Habroscelimorpha striga (LeConte, 1875)

Eunota togata fascinans (Casey, 1914)

Eunota togata globicollis (Casey, 1913)

Eunota togata togata (LaFerté-Sénectère, 1841)

Cicindela (Cicindelidia) abdominalis Fabricius, 1801

Cicindela (Cicindelidia) amargosae amargosae Dahl, 1939

Cicindela (Cicindelidia) amargosae nyensis Rumpp, 1956

Cicindela (Cicindelidia) cazieri Vogt, 1949

Cicindela (Cicindelidia) floridana Cartwright, 1939

Cicindela (Cicindelidia) hemorrhagica arizonae Wickham, 1899

Cicindela (Cicindelidia) hemorrhagica hemorrhagica LeConte, 1851

Cicindela (Cicindelidia) highlandensis Choate, 1984

Cicindela (Cicindelidia) hornii hornii Schaupp, 1883

Cicindela (Cicindelidia) marginipennis Dejean, 1831

Cicindela (Cicindelidia) nigrocoerulea bowditchi Leng, 1902

Cicindela (Cicindelidia) nigrocoerulea nigrocoerulea LeConte, 1846

Cicindela (Cicindelidia) nigrocoerulea subtropica Vogt, 1949

Cicindela (Cicindelidia) obsoleta neojuvenilis Vogt, 1949

Cicindela (Cicindelidia) obsoleta obsoleta Say, 1823

Cicindela (Cicindelidia) obsoleta santaclarae Bates, 1890

Cicindela (Cicindelidia) obsoleta vulturina LeConte, 1853

Cicindela (Cicindelidia) ocellata ocellata Klug, 1834

Cicindela (Cicindelidia) ocellata rectilatera Chaudoir, 1843

Cicindela (Cicindelidia) politula barbaraannae Sumlin, 1976

Cicindela (Cicindelidia) politula petrophila Sumlin, 1985

Cicindela (Cicindelidia) politula politula LeConte, 1875

Cicindela (Cicindelidia) politula viridimonticola Gage, 1988

Cicindela (Cicindelidia) punctulata chihuahuae Bates, 1890

Cicindela (Cicindelidia) punctulata punctulata Olivier, 1790

Cicindela (Cicindelidia) roseiventris tascosaensis Davis, 1918

Cicindela (Cicindelidia) rufiventris cumatilis LeConte, 1851

Cicindela (Cicindelidia) rufiventris hentzii Dejean, 1831

Cicindela (Cicindelidia) rufiventris rufiventris Dejean, 1825

Cicindela (Cicindelidia) scabrosa Schaupp, 1884

Cicindela (Cicindelidia) schauppii Horn, 1876

Cicindela (Cicindelidia) sedecimpunctata sedecimpunctata Klug, 1834

Cicindela (Cicindelidia) senilis Horn, 1867

Cicindela (Cicindelidia) tenuisignata LeConte, 1851

Cicindela (Cicindelidia) trifasciata ascendens LeConte, 1851

Cicindela (Cicindelidia) trifasciata sigmoidea LeConte, 1851

Cicindela (Cicindelidia) willistoni echo Casey, 1897

Cicindela (Cicindelidia) willistoni estancia Rumpp, 1962

Cicindela (Cicindelidia) willistoni funaroi Rotger, 1972

Cicindela (Cicindelidia) willistoni hirtifrons Willis, 1967

Cicindela (Cicindelidia) willistoni praedicta Rumpp, 1956

Cicindela (Cicindelidia) willistoni pseudosenilis Horn, 1900

Cicindela (Cicindelidia) willistoni sulfontis Rumpp, 1977

Cicindela (Cicindelidia) willistoni willistoni LeConte, 1879

Cicindela (Cicindela) albissima Rumpp, 1962

Cicindela (Cicindela) ancocisconensis Harris, 1852

Cicindela (Cicindela) arenicola Rumpp, 1967

Cicindela (Cicindela) arida Davis, 1928

Cicindela (Cicindela) bellissima bellissima Leng, 1902

Cicindela (Cicindela) bellissima frechini Leffler, 1979

Cicindela (Cicindela) columbica Hatch, 1938

Cicindela (Cicindela) decemnotata bonnevillensis Knisley & Kippenhan, 2012

Cicindela (Cicindela) decemnotata decemnotata Say, 1817

Cicindela (Cicindela) decemnotata meriwetheri Knisley & Kippenhan, 2012

Cicindela (Cicindela) decemnotata montevolans Knisley & Kippenhan, 2012

Cicindela (Cicindela) denikei Brown, 1934

Cicindela (Cicindela) denverensis Casey, 1897

Cicindela (Cicindela) depressula depressula Casey, 1897

Cicindela (Cicindela) depressula eureka Fall, 1901

Cicindela (Cicindela) duodecimguttata Dejean, 1825

Cicindela (Cicindela) formosa formosa Say, 1817

Cicindela (Cicindela) formosa generosa Dejean, 1831

Cicindela (Cicindela) formosa gibsoni Brown, 1940

Cicindela (Cicindela) formosa pigmentosignata Horn, 1930

Cicindela (Cicindela) formosa rutilovirescens Rumpp, 1986

Cicindela (Cicindela) fulgida fulgida Say, 1823

Cicindela (Cicindela) fulgida pseudowillistoni Horn, 1938

Cicindela (Cicindela) fulgida westbournei Calder, 1922

Cicindela (Cicindela) hirticollis abrupta Casey, 1913

Cicindela (Cicindela) hirticollis athabascensis Graves, 1988

Cicindela (Cicindela) hirticollis coloradula Graves, 1988

Cicindela (Cicindela) hirticollis corpuscula Rumpp, 1962

Cicindela (Cicindela) hirticollis couleensis Graves, 1988

Cicindela (Cicindela) hirticollis gravida LeConte, 1851

Cicindela (Cicindela) hirticollis hirticollis Say, 1817

Cicindela (Cicindela) hirticollis rhodensis Calder, 1916

Cicindela (Cicindela) hirticollis shelfordi Graves, 1988

Cicindela (Cicindela) hirticollis siuslawensis Graves, 1988

Cicindela (Cicindela) latesignata latesignata LeConte, 1851

Cicindela (Cicindela) lengi jordai Rotger, 1974

Cicindela (Cicindela) lengi lengi Horn, 1908

Cicindela (Cicindela) lengi versuta Casey, 1913

Cicindela (Cicindela) limbalis Klug, 1834

Cicindela (Cicindela) limbata hyperborea LeConte, 1863

Cicindela (Cicindela) limbata labradorensis Johnson, 1991

Cicindela (Cicindela) limbata limbata Say, 1823

Cicindela (Cicindela) limbata nogahabarensis Knisley, 2008

Cicindela (Cicindela) limbata nympha Casey, 1913

Cicindela (Cicindela) longilabris laurentii Schaupp, 1884

Cicindela (Cicindela) longilabris longilabris Say, 1824

Cicindela (Cicindela) longilabris perviridis Schaupp, 1884

Cicindela (Cicindela) nebraskana Casey, 1909

Cicindela (Cicindela) nigrior Schaupp, 1884

Cicindela (Cicindela) ohlone Freitag & Kavanaugh, 1993

Cicindela (Cicindela) oregona guttifera LeConte, 1856

Cicindela (Cicindela) oregona maricopa Leng, 1902

Cicindela (Cicindela) oregona navajoensis Van Dyke, 1947

Cicindela (Cicindela) oregona oregona LeConte, 1856

Cicindela (Cicindela) parowana parowana Wickham, 1905

Cicindela (Cicindela) parowana platti Cazier, 1937

Cicindela (Cicindela) parowana wallisi Calder, 1922

Cicindela (Cicindela) patruela consentanea Dejean, 1825

Cicindela (Cicindela) patruela patruela Dejean, 1825

Cicindela (Cicindela) pimeriana LeConte, 1867

Cicindela (Cicindela) plutonica Casey, 1897

Cicindela (Cicindela) pugetana Casey, 1914

Cicindela (Cicindela) pulchra dorothea Rumpp, 1977

Cicindela (Cicindela) pulchra pulchra Say, 1823

Cicindela (Cicindela) purpurea audubonii LeConte, 1845

Cicindela (Cicindela) purpurea cimarrona LeConte, 1868

Cicindela (Cicindela) purpurea hatchi Leffler, 1980

Cicindela (Cicindela) purpurea lauta Casey, 1897

Cicindela (Cicindela) purpurea purpurea Olivier, 1790

Cicindela (Cicindela) repanda novascotiae Vaurie, 1951

Cicindela (Cicindela) repanda repanda Dejean, 1825

Cicindela (Cicindela) repanda tanneri Knaus, 1929

Cicindela (Cicindela) scutellaris flavoviridis Vaurie, 1950

Cicindela (Cicindela) scutellaris lecontei Haldeman, 1853

Cicindela (Cicindela) scutellaris rugata Vaurie, 1950

Cicindela (Cicindela) scutellaris rugifrons Dejean, 1825

Cicindela (Cicindela) scutellaris scutellaris Say, 1823

Cicindela (Cicindela) scutellaris unicolor Dejean, 1825

Cicindela (Cicindela) scutellaris yampae Rumpp, 1986

Cicindela (Cicindela) sexguttata Fabricius, 1775

Cicindela (Cicindela) splendida Hentz, 1830

Cicindela (Cicindela) tenuicincta Schaupp, 1884

Cicindela (Cicindela) theatina Rotger, 1944

Cicindela (Cicindela) tranquebarica cibecuei Duncan, 1958

Cicindela (Cicindela) tranquebarica diffracta Casey, 1909

Cicindela (Cicindela) tranquebarica joaquinensis Knisley & Haines, 2007

Cicindela (Cicindela) tranquebarica kirbyi LeConte, 1867

Cicindela (Cicindela) tranquebarica parallelonota Casey, 1914

Cicindela (Cicindela) tranquebarica sierra Leng, 1902

Cicindela (Cicindela) tranquebarica tranquebarica Herbst, 1806

Cicindela (Cicindela) tranquebarica vibex Horn, 1867

Cicindela (Cicindela) tranquebarica viridissima Fall, 1910

Cicindela (Cicindela) waynei Leffler, 2001


Loricera (Loricera) decempunctata Eschscholtz, 1833

Loricera (Loricera) foveata LeConte, 1851

Loricera (Loricera) pilicornis congesta Mannerheim, 1853

Loricera (Loricera) pilicornis pilicornis (Fabricius, 1775)


Diacheila arctica amoena (Faldermann, 1835)

Diacheila polita (Faldermann, 1835)

Blethisa catenaria Brown, 1944

Blethisa hudsonica Casey, 1924

Blethisa julii LeConte, 1863

Blethisa multipunctata aurata Fischer von Waldheim, 1828

Blethisa oregonensis LeConte, 1853

Blethisa quadricollis Haldeman, 1847

Elaphrus (Arctelaphrus) lapponicus lapponicus Gyllenhal, 1810

Elaphrus (Arctelaphrus) lapponicus obliteratus Mannerheim, 1853

Elaphrus (Neoelaphrus) cicatricosus LeConte, 1847

Elaphrus (Neoelaphrus) clairvillei Kirby, 1837

Elaphrus (Neoelaphrus) fuliginosus Say, 1830

Elaphrus (Neoelaphrus) laevigatus LeConte, 1852

Elaphrus (Neoelaphrus) lindrothi Goulet, 1983

Elaphrus (Neoelaphrus) olivaceus LeConte, 1863

Elaphrus (Elaphrus) americanus americanus Dejean, 1831

Elaphrus (Elaphrus) americanus sylvanus Goulet, 1982

Elaphrus (Elaphrus) californicus Mannerheim, 1843

Elaphrus (Elaphrus) finitimus Casey, 1920

Elaphrus (Elaphrus) lecontei Crotch, 1876

Elaphrus (Elaphrus) marginicollis Goulet, 1983

Elaphrus (Elaphrus) mimus Goulet, 1983

Elaphrus (Elaphrus) ruscarius Say, 1830

Elaphrus (Elaphrus) trossulus Semenov, 1904

Elaphrus (Elaphrus) tuberculatus Mäklin, 1878

Elaphrus (Elaphrus) viridis Horn, 1878

Elaphrus (Elaphroterus) angusticollis angusticollis Sahlberg, 1844

Elaphrus (Elaphroterus) purpurans Hausen, 1891


Omophron (Omophron) americanum Dejean, 1831

Omophron (Omophron) dentatum LeConte, 1852

Omophron (Omophron) gilae LeConte, 1852

Omophron (Omophron) grossum Casey, 1909

Omophron (Omophron) labiatum (Fabricius, 1801)

Omophron (Omophron) nitidum LeConte, 1847

Omophron (Omophron) obliteratum Horn, 1870

Omophron (Omophron) ovale Horn, 1870

Omophron (Omophron) robustum Horn, 1870

Omophron (Omophron) solidum Casey, 1897

Omophron (Omophron) tessellatum Say, 1823


Pasimachus (Pasimachus) californicus Chaudoir, 1850

Pasimachus (Pasimachus) depressus (Fabricius, 1787)

Pasimachus (Pasimachus) duplicatus LeConte, 1853

Pasimachus (Pasimachus) elongatus LeConte, 1846

Pasimachus (Pasimachus) marginatus (Fabricius, 1787)

Pasimachus (Pasimachus) obsoletus LeConte, 1846

Pasimachus (Pasimachus) punctulatus Haldeman, 1843

Pasimachus (Pasimachus) strenuus LeConte, 1874

Pasimachus (Pasimachus) sublaevis (Palisot de Beauvois, 1811)

Pasimachus (Pasimachus) subsulcatus Say, 1823

Pasimachus (Pasimachus) viridans LeConte, 1858


Scarites (Scarites) lissopterus Chaudoir, 1881

Scarites (Scarites) marinus Nichols, 1986

Scarites (Scarites) ocalensis Nichols, 1986

Scarites (Scarites) quadriceps Chaudoir, 1843

Scarites (Scarites) stenops Bousquet & Skelley, 2010

Scarites (Scarites) subterraneus Fabricius, 1775

Scarites (Scarites) vicinus Chaudoir, 1843


Clivina (Semiclivina) dentipes Dejean, 1825

Clivina (Semiclivina) vespertina Putzeys, 1867

Clivina (Clivina) choatei Bousquet & Skelley, 2012

Clivina (Clivina) collaris (Herbst, 1784)

Clivina (Clivina) fossor fossor (Linnaeus, 1758)

Clivina (Clivina) impressefrons LeConte, 1844

Clivina (Clivina) myops Bousquet, 1997

Clivina (Clivina) oregona Fall, 1922

Clivina (Clivina) pallida Say, 1823

Clivina (Clivina) planicollis LeConte, 1857

Clivina (Clivina) punctigera LeConte, 1857

Clivina (Clivina) punctulata LeConte, 1852

Clivina (Antroforceps) alabama Bousquet, 2012

Clivina (Antroforceps) rubicunda LeConte, 1857

Clivina (Antroforceps) sasajii Ball, 2001

Clivina (Leucocara) acuducta Haldeman, 1843

Clivina (Leucocara) americana Dejean, 1831

Clivina (Leucocara) californica Van Dyke, 1925

Clivina (Leucocara) morio Dejean, 1831

Clivina (Leucocara) rufa LeConte, 1857

Paraclivina bipustulata (Fabricius, 1798)

Paraclivina convexa (LeConte, 1844)

Paraclivina fasciata (Putzeys, 1846)

Paraclivina ferrea (LeConte, 1857)

Paraclivina marginipennis (Putzeys, 1846)

Paraclivina postica (LeConte, 1846)

Paraclivina stigmula (Putzeys, 1846)

Paraclivina striatopunctata (Dejean, 1831)

Paraclivina sulcipennis (Putzeys, 1867)

Schizogenius (Genioschizus) crenulatus crenulatus LeConte, 1852

Schizogenius (Schizogenius) amphibius (Haldeman, 1843)

Schizogenius (Schizogenius) auripennis Bates, 1881

Schizogenius (Schizogenius) brevisetosus Whitehead, 1972

Schizogenius (Schizogenius) chiricahuanus Whitehead, 1972

Schizogenius (Schizogenius) depressus LeConte, 1852

Schizogenius (Schizogenius) falli Whitehead, 1972

Schizogenius (Schizogenius) ferrugineus Putzeys, 1846

Schizogenius (Schizogenius) lindrothi Whitehead, 1972

Schizogenius (Schizogenius) lineolatus (Say, 1823)

Schizogenius (Schizogenius) litigiosus Fall, 1901

Schizogenius (Schizogenius) longipennis Putzeys, 1867

Schizogenius (Schizogenius) neovalidus Whitehead, 1972

Schizogenius (Schizogenius) ochthocephalus Whitehead, 1972

Schizogenius (Schizogenius) ozarkensis Whitehead, 1972

Schizogenius (Schizogenius) planulatus LeConte, 1863

Schizogenius (Schizogenius) planuloides Whitehead, 1972

Schizogenius (Schizogenius) pluripunctatus LeConte, 1852

Schizogenius (Schizogenius) pygmaeus Van Dyke, 1925

Schizogenius (Schizogenius) sallei Putzeys, 1867

Schizogenius (Schizogenius) scopaeus Whitehead, 1972

Schizogenius (Schizogenius) seticollis seticollis Fall, 1901

Schizogenius (Schizogenius) sulcifrons Putzeys, 1846

Schizogenius (Schizogenius) tibialis Whitehead, 1972

Halocoryza arenaria (Darlington, 1939)

Oxydrepanus rufus (Putzeys, 1846)

Ardistomis obliquata Putzeys, 1846

Ardistomis schaumii LeConte, 1857

Semiardistomis puncticollis (Dejean, 1831)

Semiardistomis viridis (Say, 1823)

Aspidoglossa subangulata (Chaudoir, 1843)


Akephorus marinus LeConte, 1852

Akephorus obesus (LeConte, 1863)

Dyschirius abbreviatus Putzeys, 1846

Dyschirius aeneolus LeConte, 1850

Dyschirius affinis Fall, 1901

Dyschirius alticola Lindroth, 1961

Dyschirius analis LeConte, 1852

Dyschirius aratus LeConte, 1852

Dyschirius brevispinus LeConte, 1878

Dyschirius campicola Lindroth, 1961

Dyschirius carrorum Bousquet, 1997

Dyschirius cerberus Larson, 1968

Dyschirius chiricahuae (Dajoz, 2004)

Dyschirius comatus Bousquet, 1988

Dyschirius compactus Lindroth, 1961

Dyschirius consobrinus LeConte, 1852

Dyschirius criddlei Fall, 1925

Dyschirius curvispinus Putzeys, 1846

Dyschirius dejeanii Putzeys, 1846

Dyschirius edentulus Putzeys, 1846

Dyschirius erythrocerus LeConte, 1857

Dyschirius exochus Whitehead, 1970

Dyschirius ferrugineus Bousquet, 1988

Dyschirius gibbipennis LeConte, 1857

Dyschirius globosus (Herbst, 1784)

Dyschirius globulosus (Say, 1823)

Dyschirius haemorrhoidalis (Dejean, 1831)

Dyschirius hiemalis Bousquet, 1987

Dyschirius interior Fall, 1922

Dyschirius laevifasciatus Horn, 1878

Dyschirius larochellei Bousquet, 1988

Dyschirius longulus LeConte, 1850

Dyschirius melancholicus Putzeys, 1867

Dyschirius montanus LeConte, 1879

Dyschirius owen (Dajoz, 2004)

Dyschirius pacificus Lindroth, 1961

Dyschirius pallipennis (Say, 1823)

Dyschirius patruelis LeConte, 1852

Dyschirius perversus Fall, 1922

Dyschirius pilosus LeConte, 1857

Dyschirius planatus Lindroth, 1961

Dyschirius politus politus (Dejean, 1825)

Dyschirius pumilus (Dejean, 1825)

Dyschirius quadrimaculatus Lindroth, 1961

Dyschirius salivagans LeConte, 1875

Dyschirius sculptus Bousquet, 1988

Dyschirius sellatus LeConte, 1857

Dyschirius setosus LeConte, 1857

Dyschirius sextoni Bousquet, 1987

Dyschirius soda (Dajoz, 2004)

Dyschirius sphaericollis (Say, 1823)

Dyschirius subarcticus subarcticus Lindroth, 1961

Dyschirius sublaevis Putzeys, 1846

Dyschirius tenuispinus Lindroth, 1961

Dyschirius terminatus LeConte, 1846

Dyschirius timidus Lindroth, 1961

Dyschirius tridentatus LeConte, 1852

Dyschirius truncatus LeConte, 1857

Dyschirius unipunctatus Fall, 1901

Dyschirius varidens Fall, 1910

Dyschirius wayah (Dajoz, 2005)


Promecognathus crassus LeConte, 1868

Promecognathus laevissimus (Dejean, 1829)


Miscodera arctica (Paykull, 1798)

Broscodera (Broscodera) insignis (Mannerheim, 1852)

Zacotus matthewsii LeConte, 1869

Broscus cephalotes (Linnaeus, 1758)


Gehringia olympica Darlington, 1933


Trechoblemus westcotti Barr, 1972

Pseudanophthalmus abditus Krekeler, 1973

Pseudanophthalmus acherontis Barr, 1959

Pseudanophthalmus alabamae Valentine, 1932

Pseudanophthalmus aladdini Valentine, 1945

Pseudanophthalmus assimilis Barr, 1981

Pseudanophthalmus audax (Horn, 1883)

Pseudanophthalmus avernus Valentine, 1945

Pseudanophthalmus barberi Jeannel, 1928

Pseudanophthalmus barri Krekeler, 1973

Pseudanophthalmus beakleyi Valentine, 1937

Pseudanophthalmus bendermani Barr, 1959

Pseudanophthalmus caecus Krekeler, 1973

Pseudanophthalmus calcareus Barr, 1981

Pseudanophthalmus catherinae Barr, 1959

Pseudanophthalmus catoryctos Krekeler, 1973

Pseudanophthalmus cerberus cerberus Barr, 1985

Pseudanophthalmus cerberus completus Barr, 1985

Pseudanophthalmus chthonius Krekeler, 1973

Pseudanophthalmus ciliaris ciliaris Valentine, 1937

Pseudanophthalmus ciliaris orlindae Barr, 1959

Pseudanophthalmus cnephosus Krekeler, 1973

Pseudanophthalmus colemanensis Barr, 1959

Pseudanophthalmus conditus Krekeler, 1973

Pseudanophthalmus cordicollis Barr, 1981

Pseudanophthalmus cumberlandus Valentine, 1937

Pseudanophthalmus darlingtoni darlingtoni Barr, 1985

Pseudanophthalmus darlingtoni persimilis Barr, 1985

Pseudanophthalmus deceptivus Barr, 1981

Pseudanophthalmus delicatus Valentine, 1932

Pseudanophthalmus desertus Krekeler, 1973

Pseudanophthalmus digitus Valentine, 1932

Pseudanophthalmus distinguens Valentine, 1948

Pseudanophthalmus egberti Barr, 1965

Pseudanophthalmus elongatus Krekeler, 1973

Pseudanophthalmus emersoni Krekeler, 1958

Pseudanophthalmus engelhardti (Barber, 1928)

Pseudanophthalmus eremita (Horn, 1871)

Pseudanophthalmus exiguus Krekeler, 1973

Pseudanophthalmus exoticus Krekeler, 1973

Pseudanophthalmus farrelli Barr, 1959

Pseudanophthalmus fastigatus Barr, 1981

Pseudanophthalmus fluviatilis Valentine, 1948

Pseudanophthalmus fowlerae Barr, 1980

Pseudanophthalmus frigidus Barr, 1981

Pseudanophthalmus fulleri Valentine, 1932

Pseudanophthalmus fuscus constrictus Valentine, 1932

Pseudanophthalmus fuscus fuscus Valentine, 1931

Pseudanophthalmus georgiae Barr, 1981

Pseudanophthalmus globiceps Barr, 1985

Pseudanophthalmus gracilis Valentine, 1931

Pseudanophthalmus grandis elevatus Valentine, 1932

Pseudanophthalmus grandis grandis Valentine, 1931

Pseudanophthalmus hadenoecus Barr, 1965

Pseudanophthalmus henroti Jeannel, 1949

Pseudanophthalmus hesperus Barr, 1959

Pseudanophthalmus higginbothami Valentine, 1931

Pseudanophthalmus hirsutus Valentine, 1931

Pseudanophthalmus hoffmani Barr, 1965

Pseudanophthalmus holsingeri Barr, 1965

Pseudanophthalmus horni (Garman, 1892)

Pseudanophthalmus hortulanus Barr, 1965

Pseudanophthalmus hubbardi (Barber, 1928)

Pseudanophthalmus hubrichti Valentine, 1948

Pseudanophthalmus humeralis Valentine, 1931

Pseudanophthalmus hypertrichosis Valentine, 1932

Pseudanophthalmus hypolithos Barr, 1981

Pseudanophthalmus illinoisensis Barr & Peck, 1966

Pseudanophthalmus inexpectatus Barr, 1959

Pseudanophthalmus inquisitor Barr, 1980

Pseudanophthalmus insularis Barr, 1959

Pseudanophthalmus intermedius (Valentine, 1931)

Pseudanophthalmus intersectus Barr, 1965

Pseudanophthalmus jonesi Valentine, 1945

Pseudanophthalmus krameri Krekeler, 1973

Pseudanophthalmus krekeleri Barr, 1965

Pseudanophthalmus lallemanti Jeannel, 1949

Pseudanophthalmus leonae Barr, 1960

Pseudanophthalmus limicola Jeannel, 1931

Pseudanophthalmus lodingi Valentine, 1931

Pseudanophthalmus loganensis Barr, 1959

Pseudanophthalmus longiceps Barr, 1981

Pseudanophthalmus macradyi Valentine, 1948

Pseudanophthalmus major Krekeler, 1973

Pseudanophthalmus menetriesii campestris Barr, 1985

Pseudanophthalmus menetriesii menetriesii (Motschulsky, 1862)

Pseudanophthalmus meridionalis Valentine, 1945

Pseudanophthalmus montanus Barr, 1965

Pseudanophthalmus nelsoni Barr, 1965

Pseudanophthalmus nickajackensis Barr, 1981

Pseudanophthalmus nortoni Barr, 1981

Pseudanophthalmus occidentalis Barr, 1959

Pseudanophthalmus ohioensis Krekeler, 1973

Pseudanophthalmus orientalis Krekeler, 1973

Pseudanophthalmus orthosulcatus Valentine, 1932

Pseudanophthalmus packardi Barr, 1959

Pseudanophthalmus pallidus Barr, 1981

Pseudanophthalmus paradoxus Barr, 1981

Pseudanophthalmus parvicollis Jeannel, 1931

Pseudanophthalmus parvus Krekeler, 1973

Pseudanophthalmus paulus Barr, 1981

Pseudanophthalmus paynei Barr, 1981

Pseudanophthalmus petrunkevitchi Valentine, 1945

Pseudanophthalmus pholeter Krekeler, 1973

Pseudanophthalmus pilosus Barr, 1985

Pseudanophthalmus pontis Barr, 1965

Pseudanophthalmus potomaca Valentine, 1932

Pseudanophthalmus praetermissus Barr, 1981

Pseudanophthalmus princeps Barr, 1979

Pseudanophthalmus productus Barr, 1980

Pseudanophthalmus profundus Valentine, 1945

Pseudanophthalmus pubescens intrepidus Barr, 1985

Pseudanophthalmus pubescens pubescens (Horn, 1869)

Pseudanophthalmus punctatus Valentine, 1931

Pseudanophthalmus pusillus Barr, 1981

Pseudanophthalmus pusio (Horn, 1869)

Pseudanophthalmus puteanus Krekeler, 1973

Pseudanophthalmus quadratus Barr, 1965

Pseudanophthalmus rittmani Krekeler, 1973

Pseudanophthalmus robustus Valentine, 1931

Pseudanophthalmus rogersae Barr, 1981

Pseudanophthalmus rotundatus Valentine, 1932

Pseudanophthalmus sanctipauli Barr, 1981

Pseudanophthalmus scholasticus Barr, 1981

Pseudanophthalmus scutilus Barr, 1981

Pseudanophthalmus seclusus Barr, 1981

Pseudanophthalmus senecae Valentine, 1932

Pseudanophthalmus sequoyah Barr, 1981

Pseudanophthalmus sericus Barr, 1981

Pseudanophthalmus shilohensis mayfieldensis Krekeler, 1958

Pseudanophthalmus shilohensis shilohensis Krekeler, 1958

Pseudanophthalmus sidus Barr, 1965

Pseudanophthalmus simplex Barr, 1980

Pseudanophthalmus simulans Barr, 1985

Pseudanophthalmus solivagus Krekeler, 1973

Pseudanophthalmus steevesi Barr, 1981

Pseudanophthalmus striatus (Motschulsky, 1862)

Pseudanophthalmus stricticollis Jeannel, 1931

Pseudanophthalmus sylvaticus Barr, 1967

Pseudanophthalmus templetoni Valentine, 1948

Pseudanophthalmus tenebrosus Krekeler, 1973

Pseudanophthalmus tennesseensis Valentine, 1937

Pseudanophthalmus tenuis (Horn, 1871)

Pseudanophthalmus thomasi Barr, 1981

Pseudanophthalmus tiresias Barr, 1959

Pseudanophthalmus transfluvialis Barr, 1985

Pseudanophthalmus troglodytes Krekeler, 1973

Pseudanophthalmus tullahoma Barr, 1959

Pseudanophthalmus umbratilis Krekeler, 1973

Pseudanophthalmus unionis Barr, 1981

Pseudanophthalmus valentinei Jeannel, 1949

Pseudanophthalmus vanburenensis Barr, 1959

Pseudanophthalmus ventus Barr, 1981

Pseudanophthalmus vicarius Barr, 1965

Pseudanophthalmus virginicus (Barr, 1960)

Pseudanophthalmus wallacei Barr, 1981

Pseudanophthalmus youngi Krekeler, 1958

Nelsonites jonesei Valentine, 1952

Nelsonites walteri Valentine, 1952

Neaphaenops tellkampfii henroti Jeannel, 1949

Neaphaenops tellkampfii meridionalis Barr, 1959

Neaphaenops tellkampfii tellkampfii (Erichson, 1844)

Neaphaenops tellkampfii viator Barr, 1979

Blemus discus discus (Fabricius, 1792)

Xenotrechus condei Barr & Krekeler, 1967

Xenotrechus denticollis Barr & Krekeler, 1967

Darlingtonea kentuckensis Valentine, 1952

Ameroduvalius jeanneli jeanneli Valentine, 1952

Ameroduvalius jeanneli rockcastlei Valentine, 1952

Trechus (Trechus) alinae Dajoz, 1990

Trechus (Trechus) apache Dajoz, 1990

Trechus (Trechus) apicalis Motschulsky, 1845

Trechus (Trechus) arizonae Casey, 1918

Trechus (Trechus) caliginis Barr, 1985

Trechus (Trechus) carolinae Schaeffer, 1901

Trechus (Trechus) chalybeus Dejean, 1831

Trechus (Trechus) coloradensis Schaeffer, 1915

Trechus (Trechus) conformis Jeannel, 1927

Trechus (Trechus) crassiscapus Lindroth, 1955

Trechus (Trechus) cumberlandus Barr, 1962

Trechus (Trechus) humboldti Van Dyke, 1945

Trechus (Trechus) hydropicus avus Barr, 1962

Trechus (Trechus) hydropicus beutenmuelleri Jeannel, 1931

Trechus (Trechus) hydropicus canus Barr, 1962

Trechus (Trechus) hydropicus hydropicus Horn, 1883

Trechus (Trechus) mitchellensis Barr, 1962

Trechus (Trechus) obtusus Erichson, 1837

Trechus (Trechus) oregonensis Hatch, 1951

Trechus (Trechus) ovipennis Motschulsky, 1845

Trechus (Trechus) pomonae Fall, 1901

Trechus (Trechus) quadristriatus (Schrank, 1781)

Trechus (Trechus) roanicus Barr, 1962

Trechus (Trechus) rubens (Fabricius, 1792)

Trechus (Trechus) schwarzi saludae Barr, 1979

Trechus (Trechus) schwarzi schwarzi Jeannel, 1931

Trechus (Trechus) schwarzi scopulosus Barr, 1979

Trechus (Trechus) tenuiscapus Lindroth, 1961

Trechus (Trechus) yvesbousqueti Donabauer, 2010

Trechus (Microtrechus) aduncus Barr, 1962

Trechus (Microtrechus) balsamensis Barr, 1962

Trechus (Microtrechus) barberi (Jeannel, 1931)

Trechus (Microtrechus) bowlingi Barr, 1962

Trechus (Microtrechus) cheoahensis Donabauer, 2005

Trechus (Microtrechus) clingmanensis Donabauer, 2005

Trechus (Microtrechus) coweensis Barr, 1979

Trechus (Microtrechus) haoe Barr, 1979

Trechus (Microtrechus) haoeleadensis Donabauer, 2005

Trechus (Microtrechus) howellae Barr, 1979

Trechus (Microtrechus) inexpectatus Barr, 1985

Trechus (Microtrechus) luculentus cheoahbaldensis Donabauer, 2005

Trechus (Microtrechus) luculentus joannabaldensis Donabauer, 2005

Trechus (Microtrechus) luculentus luculentus Barr, 1962

Trechus (Microtrechus) luculentus wayahensis Barr, 1979

Trechus (Microtrechus) nantahalae Barr, 1979

Trechus (Microtrechus) nebulosus Barr, 1962

Trechus (Microtrechus) novaculosus Barr, 1962

Trechus (Microtrechus) plottbalsamensis Donabauer, 2005

Trechus (Microtrechus) pseudobarberi Donabauer, 2009

Trechus (Microtrechus) pseudonovaculosus Donabauer, 2005

Trechus (Microtrechus) pseudosubtilis Donabauer, 2009

Trechus (Microtrechus) ramseyensis Donabauer, 2005

Trechus (Microtrechus) rivulis Dajoz, 2005

Trechus (Microtrechus) rosenbergi Barr, 1962

Trechus (Microtrechus) satanicus Barr, 1962

Trechus (Microtrechus) snowbirdensis Donabauer, 2005

Trechus (Microtrechus) stefanschoedli Donabauer, 2005

Trechus (Microtrechus) stupkai Barr, 1979

Trechus (Microtrechus) subtilis Barr, 1962

Trechus (Microtrechus) talequah Barr, 1962

Trechus (Microtrechus) tennesseensis tauricus Barr, 1962

Trechus (Microtrechus) tennesseensis tennesseensis Barr, 1962

Trechus (Microtrechus) thomasbarri Donabauer, 2005

Trechus (Microtrechus) thunderheadensis Donabauer, 2005

Trechus (Microtrechus) tobiasi Donabauer, 2005

Trechus (Microtrechus) tonitru Barr, 1962

Trechus (Microtrechus) toxawayi Barr, 1979

Trechus (Microtrechus) tuckaleechee Barr, 1962

Trechus (Microtrechus) tusquitee Barr, 1979

Trechus (Microtrechus) tusquitensis Donabauer, 2005

Trechus (Microtrechus) uncifer Barr, 1962

Trechus (Microtrechus) unicoi Barr, 1979

Trechus (Microtrechus) valentinei Barr, 1979

Trechus (Microtrechus) vandykei pisgahensis Barr, 1979

Trechus (Microtrechus) vandykei vandykei (Jeannel, 1927)

Trechus (Microtrechus) verus Barr, 1962

Trechus (Microtrechus) wayahbaldensis Donabauer, 2005


Amerizus (Amerizus) oblonguloides (Lindroth, 1963)

Amerizus (Amerizus) oblongulus (Mannerheim, 1852)

Amerizus (Amerizus) spectabilis (Mannerheim, 1852)

Amerizus (Amerizus) utahensis (Van Dyke, 1926)

Amerizus (Amerizus) wingatei (Bland, 1864)

Lionepha casta (Casey, 1918)

Lionepha chintimini (Erwin & Kavanaugh, 1981)

Lionepha disjuncta (Lindroth, 1963)

Lionepha erasa (LeConte, 1859)

Lionepha lindrothellus (Erwin & Kavanaugh, 1981)

Lionepha lummi (Erwin & Kavanaugh, 1981)

Lionepha osculans (Casey, 1918)

Lionepha pseudoerasa (Lindroth, 1963)

Lionepha sequoiae (Lindroth, 1963)

Asaphidion alaskanum Wickham, 1919

Asaphidion curtum curtum (Heyden, 1870)

Asaphidion yukonense Wickham, 1919

Bembidion (Hirmoplataphus) alpineanum Casey, 1924

Bembidion (Hirmoplataphus) avidum Casey, 1918

Bembidion (Hirmoplataphus) concolor (Kirby, 1837)

Bembidion (Hirmoplataphus) humboldtense Blaisdell, 1902

Bembidion (Hirmoplataphus) nigrum Say, 1823

Bembidion (Hirmoplataphus) quadrulum LeConte, 1861

Bembidion (Hirmoplataphus) recticolle LeConte, 1863

Bembidion (Hirmoplataphus) salebratum (LeConte, 1847)

Bembidion (Hirmoplataphus) subaerarium Casey, 1924

Bembidion (Hydriomicrus) brevistriatum Hayward, 1897

Bembidion (Hydriomicrus) californicum Hayward, 1897

Bembidion (Hydriomicrus) innocuum Casey, 1918

Bembidion (Hydriomicrus) quadratulum Notman, 1920

Bembidion (Hydriomicrus) semistriatum (Haldeman, 1843)

Bembidion (Odontium) aenulum Hayward, 1901

Bembidion (Odontium) bowditchii LeConte, 1878

Bembidion (Odontium) carinatum (LeConte, 1852)

Bembidion (Odontium) confusum Hayward, 1897

Bembidion (Odontium) coxendix Say, 1823

Bembidion (Odontium) durangoense Bates, 1891

Bembidion (Odontium) gilae Lindroth, 1963

Bembidion (Odontium) paraenulum Maddison, 2009

Bembidion (Odontium) robusticolle Hayward, 1897

Bembidion (Odontium) sculpturatum (Motschulsky, 1859)

Bembidion (Bracteon) alaskense Lindroth, 1962

Bembidion (Bracteon) balli Lindroth, 1962

Bembidion (Bracteon) carinula Chaudoir, 1868

Bembidion (Bracteon) foveum Motschulsky, 1844

Bembidion (Bracteon) hesperium Fall, 1910

Bembidion (Bracteon) inaequale Say, 1823

Bembidion (Bracteon) lapponicum Zetterstedt, 1828

Bembidion (Bracteon) levettei carrianum Casey, 1924

Bembidion (Bracteon) levettei levettei Casey, 1918

Bembidion (Bracteon) lorquinii Chaudoir, 1868

Bembidion (Bracteon) punctatostriatum Say, 1823

Bembidion (Bracteon) zephyrum Fall, 1910

Bembidion (Ochthedromus) americanum Dejean, 1831

Bembidion (Ochthedromus) bifossulatum (LeConte, 1852)

Bembidion (Ochthedromus) cheyennense Casey, 1918

Bembidion (Pseudoperyphus) antiquum Dejean, 1831

Bembidion (Pseudoperyphus) arenobile Maddison, 2008

Bembidion (Pseudoperyphus) bellorum Maddison, 2008

Bembidion (Pseudoperyphus) chalceum Dejean, 1831

Bembidion (Pseudoperyphus) honestum Say, 1823

Bembidion (Pseudoperyphus) integrum Casey, 1918

Bembidion (Pseudoperyphus) louisella Maddison, 2008

Bembidion (Pseudoperyphus) rothfelsi Maddison, 2008

Bembidion (Pseudoperyphus) rufotinctum Chaudoir, 1868

Bembidion (Cillenus) palosverdes Kavanaugh & Erwin, 1992

Bembidion (Actedium) lachnophoroides Darlington, 1926

Bembidion (Ocydromus) scopulinum (Kirby, 1837)

Bembidion (Peryphus) actuosum Casey, 1918

Bembidion (Peryphus) bruxellense Wesmael, 1835

Bembidion (Peryphus) consanguineum Hayward, 1897

Bembidion (Peryphus) dauricum (Motschulsky, 1844)

Bembidion (Peryphus) femoratum femoratum Sturm, 1825

Bembidion (Peryphus) lugubre LeConte, 1857

Bembidion (Peryphus) mexicanum Dejean, 1831

Bembidion (Peryphus) nevadense Ulke, 1875

Bembidion (Peryphus) obscurellum obscurellum (Motschulsky, 1845)

Bembidion (Peryphus) pernotum Casey, 1918

Bembidion (Peryphus) perspicuum (LeConte, 1848)

Bembidion (Peryphus) petrosum attuense Lindroth, 1963

Bembidion (Peryphus) petrosum petrosum Gebler, 1833

Bembidion (Peryphus) plagiatum (Zimmermann, 1869)

Bembidion (Peryphus) poppii schalleri Lindroth, 1963

Bembidion (Peryphus) rupicola (Kirby, 1837)

Bembidion (Peryphus) sarpedon Casey, 1918

Bembidion (Peryphus) satelles Casey, 1918

Bembidion (Peryphus) sejunctum sejunctum Casey, 1918

Bembidion (Peryphus) sejunctum semiaureum Fall, 1922

Bembidion (Peryphus) striola (LeConte, 1852)

Bembidion (Peryphus) tetracolum tetracolum Say, 1823

Bembidion (Peryphus) transversale Dejean, 1831

Bembidion (Terminophanes) mckinleyi carneum Lindroth, 1963

Bembidion (Terminophanes) mckinleyi mckinleyi Fall, 1926

Bembidion (Asioperyphus) bimaculatum (Kirby, 1837)

Bembidion (Asioperyphus) lenae Csiki, 1928

Bembidion (Asioperyphus) postremum Say, 1830

Bembidion (Asioperyphus) renoanum Casey, 1918

Bembidion (Asioperyphus) sordidum (Kirby, 1837)

Bembidion (Asioperyphus) umiatense Lindroth, 1963

Bembidion (Peryphanes) grapii Gyllenhal, 1827

Bembidion (Peryphanes) lacunarium (Zimmermann, 1869)

Bembidion (Peryphanes) platynoides Hayward, 1897

Bembidion (Peryphanes) stephensii Crotch, 1866

Bembidion (Peryphanes) subangustatum Hayward, 1897

Bembidion (Peryphanes) texanum Chaudoir, 1868

Bembidion (Peryphanes) yukonum Fall, 1926

Bembidion (Testediolum) commotum Casey, 1918

Bembidion (Testediolum) modocianum Casey, 1924

Bembidion (Testediolum) nebraskense LeConte, 1863

Bembidion (Testediolum) obscuripenne Blaisdell, 1902

Bembidion (Testediolum) perbrevicolle Casey, 1924

Bembidion (Testediolum) ulkei Lindroth, 1963

Bembidion (Leuchydrium) tigrinum LeConte, 1879

Bembidion (Bembidion) adductum Casey, 1918

Bembidion (Bembidion) mutatum Gemminger & Harold, 1868

Bembidion (Bembidion) oregonense Hatch, 1953

Bembidion (Bembidion) pedicellatum LeConte, 1857

Bembidion (Bembidion) praecinctum LeConte, 1879

Bembidion (Bembidion) quadrimaculatum dubitans (LeConte, 1852)

Bembidion (Bembidion) quadrimaculatum oppositum Say, 1823

Bembidion (Cyclolopha) jucundum Horn, 1895

Bembidion (Cyclolopha) poculare Bates, 1884

Bembidion (Cyclolopha) sphaeroderum Bates, 1882

Bembidion (Furcacampa) affine Say, 1823

Bembidion (Furcacampa) egens Casey, 1918

Bembidion (Furcacampa) fuchsii Blaisdell, 1902

Bembidion (Furcacampa) impotens Casey, 1918

Bembidion (Furcacampa) mimus Hayward, 1897

Bembidion (Furcacampa) nogalesium Casey, 1924

Bembidion (Furcacampa) timidum (LeConte, 1847)

Bembidion (Furcacampa) triviale Casey, 1918

Bembidion (Furcacampa) versicolor (LeConte, 1847)

Bembidion (Neobembidion) constricticolle Hayward, 1897

Bembidion (Neobembidion) nitidicolle Bousquet, 2006

Bembidion (Neobembidion) nudipenne Lindroth, 1963

Bembidion (Neobembidion) tencenti Hatch, 1951

Bembidion (Diplocampa) transparens transparens (Gebler, 1830)

Bembidion (Semicampa) convexulum Hayward, 1897

Bembidion (Semicampa) morulum LeConte, 1863

Bembidion (Semicampa) muscicola Hayward, 1897

Bembidion (Semicampa) nigrivestis Bousquet, 2006

Bembidion (Semicampa) praticola Lindroth, 1963

Bembidion (Semicampa) roosevelti Pic, 1902

Bembidion (Semicampa) rubiginosum LeConte, 1879

Bembidion (Semicampa) semicinctum Notman, 1919

Bembidion (Notaphus) acticola Casey, 1884

Bembidion (Notaphus) aeneicolle (LeConte, 1847)

Bembidion (Notaphus) approximatum (LeConte, 1852)

Bembidion (Notaphus) aratum (LeConte, 1852)

Bembidion (Notaphus) auxiliator Casey, 1924

Bembidion (Notaphus) callens Casey, 1918

Bembidion (Notaphus) castor Lindroth, 1963

Bembidion (Notaphus) coloradense Hayward, 1897

Bembidion (Notaphus) consimile Hayward, 1897

Bembidion (Notaphus) conspersum Chaudoir, 1868

Bembidion (Notaphus) constrictum (Leconte, 1847)

Bembidion (Notaphus) consuetum Casey, 1918

Bembidion (Notaphus) contractum Say, 1823

Bembidion (Notaphus) cordatum (LeConte, 1847)

Bembidion (Notaphus) debiliceps Casey, 1918

Bembidion (Notaphus) dejectum Casey, 1884

Bembidion (Notaphus) dorsale Say, 1823

Bembidion (Notaphus) evidens Casey, 1918

Bembidion (Notaphus) flohri Bates, 1878

Bembidion (Notaphus) graphicum Casey, 1918

Bembidion (Notaphus) hageni Hayward, 1897

Bembidion (Notaphus) idoneum Casey, 1918

Bembidion (Notaphus) indistinctum Dejean, 1831

Bembidion (Notaphus) insulatum (LeConte, 1852)

Bembidion (Notaphus) intermedium (Kirby, 1837)

Bembidion (Notaphus) jacobianum Casey, 1918

Bembidion (Notaphus) latebricola Casey, 1918

Bembidion (Notaphus) lecontei Csiki, 1928

Bembidion (Notaphus) luculentum Casey, 1918

Bembidion (Notaphus) mormon Hayward, 1897

Bembidion (Notaphus) nigripes (Kirby, 1837)

Bembidion (Notaphus) nubiculosum Chaudoir, 1868

Bembidion (Notaphus) oberthueri Hayward, 1901

Bembidion (Notaphus) obscuromaculatum (Motschulsky, 1859)

Bembidion (Notaphus) obtusangulum LeConte, 1863

Bembidion (Notaphus) obtusidens Fall, 1922

Bembidion (Notaphus) operosum Casey, 1918

Bembidion (Notaphus) patruele Dejean, 1831

Bembidion (Notaphus) pilatei Chaudoir, 1868

Bembidion (Notaphus) pimanum Casey, 1918

Bembidion (Notaphus) rapidum (LeConte, 1847)

Bembidion (Notaphus) scintillans Bates, 1882

Bembidion (Notaphus) scudderi LeConte, 1878

Bembidion (Notaphus) semiopacum Casey, 1924

Bembidion (Notaphus) semipunctatum (Donovan, 1806)

Bembidion (Notaphus) umbratum (LeConte, 1847)

Bembidion (Notaphus) versutum LeConte, 1878

Bembidion (Notaphus) viridicolle (LaFerté-Sénectère, 1841)

Bembidion (Notaphus) vividum Casey, 1884

Bembidion (Notaphus) vulpecula Casey, 1918

Bembidion (Trepanedoris) acutifrons LeConte, 1879

Bembidion (Trepanedoris) ampliceps Casey, 1918

Bembidion (Trepanedoris) anguliferum (LeConte, 1852)

Bembidion (Trepanedoris) canadianum Casey, 1924

Bembidion (Trepanedoris) clemens Casey, 1918

Bembidion (Trepanedoris) concretum Casey, 1918

Bembidion (Trepanedoris) connivens (LeConte, 1852)

Bembidion (Trepanedoris) elizabethae Hatch, 1950

Bembidion (Trepanedoris) fortestriatum (Motschulsky, 1845)

Bembidion (Trepanedoris) frontale (LeConte, 1847)

Bembidion (Trepanedoris) pseudocautum Lindroth, 1963

Bembidion (Trepanedoris) scenicum Casey, 1918

Bembidion (Trepanedoris) siticum Casey, 1918

Bembidion (Peryphodes) ephippigerum (LeConte, 1852)

Bembidion (Peryphodes) salinarium Casey, 1918

Bembidion (Emphanes) diligens Casey, 1918

Bembidion (Emphanes) vile (LeConte, 1852)

Bembidion (Blepharoplataphus) hastii Sahlberg, 1827

Bembidion (Plataphus) arcticum Lindroth, 1963

Bembidion (Plataphus) basicorne Notman, 1920

Bembidion (Plataphus) brachythorax Lindroth, 1963

Bembidion (Plataphus) breve (Motschulsky, 1845)

Bembidion (Plataphus) carolinense Casey, 1924

Bembidion (Plataphus) complanulum (Mannerheim, 1853)

Bembidion (Plataphus) compressum Lindroth, 1963

Bembidion (Plataphus) curtulatum Casey, 1918

Bembidion (Plataphus) falsum Blaisdell, 1902

Bembidion (Plataphus) farrarae Hatch, 1950

Bembidion (Plataphus) gebleri turbatum Casey, 1918

Bembidion (Plataphus) gordoni Lindroth, 1963

Bembidion (Plataphus) gratiosum Casey, 1918

Bembidion (Plataphus) haruspex Casey, 1918

Bembidion (Plataphus) hyperboraeorum Munster, 1923

Bembidion (Plataphus) improvidens Casey, 1924

Bembidion (Plataphus) kalumae Lindroth, 1963

Bembidion (Plataphus) kuprianovii Mannerheim, 1843

Bembidion (Plataphus) laxatum Casey, 1918

Bembidion (Plataphus) manningense Lindroth, 1969

Bembidion (Plataphus) neocoerulescens Bousquet, 1993

Bembidion (Plataphus) nigrocoeruleum Hayward, 1897

Bembidion (Plataphus) occultator Notman, 1920

Bembidion (Plataphus) oppressum Casey, 1918

Bembidion (Plataphus) placeranum Casey, 1924

Bembidion (Plataphus) planiusculum Mannerheim, 1843

Bembidion (Plataphus) quadrifoveolatum Mannerheim, 1843

Bembidion (Plataphus) rosslandicum Lindroth, 1963

Bembidion (Plataphus) rufinum Lindroth, 1963

Bembidion (Plataphus) rusticum lenensoides Lindroth, 1963

Bembidion (Plataphus) rusticum rusticum Casey, 1918

Bembidion (Plataphus) sierricola Casey, 1924

Bembidion (Plataphus) simplex Hayward, 1897

Bembidion (Plataphus) stillaguamish Hatch, 1950

Bembidion (Plataphus) sulcipenne hyperboroides Lindroth, 1963

Bembidion (Plataphus) sulcipenne prasinoides Lindroth, 1963

Bembidion (Plataphus) vandykei Blaisdell, 1902

Bembidion (Plataphus) viator Casey, 1918

Bembidion (Hydrium) interventor Lindroth, 1963

Bembidion (Hydrium) levigatum Say, 1823

Bembidion (Hydrium) nitidum (Kirby, 1837)

Bembidion (Hydrium) obliquulum LeConte, 1859

Bembidion (Metallina) dyschirinum LeConte, 1861

Bembidion (Metallina) lampros (Herbst, 1784)

Bembidion (Metallina) properans (Stephens, 1828)

Bembidion (Lindrochthus) wickhami Hayward, 1897

Bembidion (Eupetedromus) graciliforme Hayward, 1897

Bembidion (Eupetedromus) immaturum Lindroth, 1954

Bembidion (Eupetedromus) incrematum LeConte, 1860

Bembidion (Eupetedromus) iridipenne Bousquet & Webster, 2006

Bembidion (Eupetedromus) variegatum Say, 1823

Bembidion (Trechonepha) iridescens (LeConte, 1852)

Bembidion (Trechonepha) trechiforme (LeConte, 1852)

Bembidion (Liocosmius) festivum Casey, 1918

Bembidion (Liocosmius) horni Hayward, 1897

Bembidion (Liocosmius) mundum (LeConte, 1852)

Bembidion (Melomalus) planatum (LeConte, 1847)

Bembidion (Trichoplataphus) fugax (LeConte, 1848)

Bembidion (Trichoplataphus) grandiceps Hayward, 1897

Bembidion (Trichoplataphus) ozarkense Maddison & Hildebrandt, 2011

Bembidion (Trichoplataphus) planum (Haldeman, 1843)

Bembidion (Trichoplataphus) rolandi Fall, 1922

Bembidion (Phyla) obtusum Audinet-Serville, 1821

Bembidion (Lymnaeum) laticeps (LeConte, 1858)

Bembidion (Lymnaeum) nigropiceum (Marsham, 1802)

Phrypeus rickseckeri (Hayward, 1897)

Mioptachys flavicauda (Say, 1823)

Tachyta (Tachyta) angulata Casey, 1918

Tachyta (Tachyta) falli (Hayward, 1900)

Tachyta (Tachyta) inornata (Say, 1823)

Tachyta (Tachyta) kirbyi Casey, 1918

Tachyta (Tachyta) parvicornis Notman, 1922

Elaphropus (Barytachys) anceps (LeConte, 1848)

Elaphropus (Barytachys) anthrax (LeConte, 1852)

Elaphropus (Barytachys) brevis (Casey, 1918)

Elaphropus (Barytachys) brunnicollis (Motschulsky, 1862)

Elaphropus (Barytachys) capax (LeConte, 1863)

Elaphropus (Barytachys) cockerelli (Fall, 1907)

Elaphropus (Barytachys) congener (Casey, 1918)

Elaphropus (Barytachys) conjugens (Notman, 1919)

Elaphropus (Barytachys) dolosus (LeConte, 1848)

Elaphropus (Barytachys) fatuus (Casey, 1918)

Elaphropus (Barytachys) ferrugineus (Dejean, 1831)

Elaphropus (Barytachys) fuscicornis (Chaudoir, 1868)

Elaphropus (Barytachys) granarius (Dejean, 1831)

Elaphropus (Barytachys) incurvus (Say, 1830)

Elaphropus (Barytachys) liebecki (Hayward, 1900)

Elaphropus (Barytachys) monticola (Casey, 1918)

Elaphropus (Barytachys) nebulosus (Chaudoir, 1868)

Elaphropus (Barytachys) obesulus (LeConte, 1852)

Elaphropus (Barytachys) rapax (LeConte, 1852)

Elaphropus (Barytachys) renoicus (Casey, 1918)

Elaphropus (Barytachys) rubricauda (Casey, 1918)

Elaphropus (Barytachys) saturatus (Casey, 1918)

Elaphropus (Barytachys) sectator (Casey, 1918)

Elaphropus (Barytachys) sedulus (Casey, 1918)

Elaphropus (Barytachys) tahoensis (Casey, 1918)

Elaphropus (Barytachys) tripunctatus (Say, 1830)

Elaphropus (Barytachys) vernicatus (Casey, 1918)

Elaphropus (Barytachys) vivax (LeConte, 1848)

Elaphropus (Barytachys) xanthopus (Dejean, 1831)

Elaphropus (Tachyura) parvulus (Dejean, 1831)

Micratopus aenescens (LeConte, 1848)

Pericompsus (Pericompsus) ephippiatus (Say, 1830)

Pericompsus (Pericompsus) laetulus LeConte, 1852

Pericompsus (Pericompsus) sellatus LeConte, 1852

Porotachys bisulcatus (Nicolai, 1822)

Polyderis diaphana (Casey, 1918)

Polyderis laeva (Say, 1823)

Polyderis rufotestacea (Hayward, 1900)

Tachys (Tachys) bradycellinus Hayward, 1900

Tachys (Tachys) bryanti Lindroth, 1966

Tachys (Tachys) corax LeConte, 1852

Tachys (Tachys) halophilus Lindroth, 1966

Tachys (Tachys) litoralis Casey, 1884

Tachys (Tachys) misellus LaFerté-Sénectère, 1841

Tachys (Tachys) mordax LeConte, 1852

Tachys (Tachys) pallidus Chaudoir, 1868

Tachys (Tachys) pulchellus LaFerté-Sénectère, 1841

Tachys (Tachys) translucens Darlington, 1937

Tachys (Tachys) virgo LeConte, 1852

Tachys (Tachys) vittiger LeConte, 1852

Tachys (Paratachys) aeneipennis Motschulsky, 1862

Tachys (Paratachys) albipes LeConte, 1863

Tachys (Paratachys) austinicus (Casey, 1918)

Tachys (Paratachys) columbiensis Hayward, 1900

Tachys (Paratachys) edax LeConte, 1852

Tachys (Paratachys) hyalinus Casey, 1918

Tachys (Paratachys) oblitus Casey, 1918

Tachys (Paratachys) potomaca (Erwin, 1981)

Tachys (Paratachys) proximus (Say, 1823)

Tachys (Paratachys) pumilus (Dejean, 1831)

Tachys (Paratachys) rectangulus Notman, 1919

Tachys (Paratachys) rhodeanus Casey, 1918

Tachys (Paratachys) sagax Casey, 1918

Tachys (Paratachys) scitulus LeConte, 1848

Tachys (Paratachys) sequax LeConte, 1848

Tachys (Paratachys) spadix Casey, 1918

Tachys (Paratachys) umbripennis Chaudoir, 1868

Tachys (Paratachys) ventricosus LeConte, 1863

Tachys (Paratachys) vernilis Casey, 1918

Tachys (Paratachys) vorax LeConte, 1852

Anillodes debilis (LeConte, 1853)

Anillodes minutus Jeannel, 1963

Anillodes walkeri Jeannel, 1963

Anillinus affabilis (Brues, 1902)

Anillinus aleyae Sokolov & Watrous, 2008

Anillinus balli Sokolov & Carlton, 2004

Anillinus barberi Jeannel, 1963

Anillinus barri Sokolov & Carlton, 2004

Anillinus campbelli Giachino, 2011

Anillinus carltoni Sokolov, 2011

Anillinus chandleri Sokolov, 2011

Anillinus cherokee Sokolov & Carlton, 2008

Anillinus chilhowee Sokolov, 2011

Anillinus cieglerae Sokolov & Carlton, 2007

Anillinus cornelli Sokolov & Carlton, 2004

Anillinus daggyi Sokolov & Carlton, 2004

Anillinus depressus (Jeannel, 1963)

Anillinus docwatsoni Sokolov & Carlton, 2004

Anillinus dohrni (Ehlers, 1884)

Anillinus elongatus Jeannel, 1963

Anillinus erwini Sokolov & Carlton, 2004

Anillinus folkertsi Sokolov & Carlton, 2004

Anillinus fortis (Horn, 1869)

Anillinus gimmeli Sokolov & Carlton, 2010

Anillinus indianae Jeannel, 1963

Anillinus juliae Sokolov & Carlton, 2010

Anillinus kovariki Sokolov & Carlton, 2004

Anillinus langdoni Sokolov & Carlton, 2004

Anillinus lescheni Sokolov & Carlton, 2004

Anillinus longiceps Jeannel, 1963

Anillinus loweae Sokolov & Carlton, 2004

Anillinus magazinensis Sokolov & Carlton, 2004

Anillinus merritti Sokolov & Carlton, 2010

Anillinus moseleyae Sokolov & Carlton, 2004

Anillinus murrayae Sokolov & Carlton, 2004

Anillinus nantahala Dajoz, 2005

Anillinus pecki Giachino, 2011

Anillinus pusillus Sokolov & Carlton, 2007

Anillinus robisoni Sokolov & Carlton, 2004

Anillinus sinuaticollis Jeannel, 1963

Anillinus sinuatus Jeannel, 1963

Anillinus smokiensis Sokolov, 2011

Anillinus steevesi Barr, 1995

Anillinus stephani Sokolov & Carlton, 2004

Anillinus tishechkini Sokolov & Carlton, 2004

Anillinus turneri Jeannel, 1963

Anillinus unicoi Sokolov, 2011

Anillinus valentinei (Jeannel, 1963)

Anillinus virginiae Jeannel, 1963

Serranillus dunavani (Jeannel, 1963)

Serranillus jeanneli Barr, 1995

Serranillus septentrionis Sokolov & Carlton, 2008

Anillaspis caseyi Jeannel, 1963

Anillaspis explanata (Horn, 1888)

Horologion speokoites Valentine, 1932


Thalassotrechus barbarae (Horn, 1892)

Diplochaetus emaciatus (Bates, 1891)

Diplochaetus megacephalus Bousquet & Laplante, 1997

Diplochaetus planatus (Horn, 1876)

Diplochaetus rutilus (Chevrolat, 1863)

Pogonus texanus Chaudoir, 1868


Diplous (Platidius) aterrimus (Dejean, 1828)

Diplous (Platidius) californicus (Motschulsky, 1844)

Diplous (Platidius) filicornis (Casey, 1918)

Diplous (Platidius) rugicollis (Randall, 1838)

Patrobus cinctus Motschulsky, 1860

Patrobus fossifrons (Eschscholtz, 1823)

Patrobus foveocollis (Eschscholtz, 1823)

Patrobus lecontei Chaudoir, 1872

Patrobus longicornis (Say, 1823)

Patrobus septentrionis septentrionis Dejean, 1828

Patrobus stygicus Chaudoir, 1872

Platypatrobus lacustris Darlington, 1938

Platidiolus vandykei Kurnakov, 1960


Nomius pygmaeus (Dejean, 1831)

Psydrus piceus LeConte, 1846


Metrius contractus contractus Eschscholtz, 1829

Metrius contractus planatus Van Dyke, 1925

Metrius contractus sericeus Rivers, 1900

Metrius explodens Bousquet & Goulet, 1990


Pachyteles gyllenhalii (Dejean, 1825)

Goniotropis kuntzeni kuntzeni Bänninger, 1927

Goniotropis parca (LeConte, 1884)

Physea hirta LeConte, 1853

Ozaena lemoulti Bänninger, 1932


Brachinus (Neobrachinus) aabaaba Erwin, 1970

Brachinus (Neobrachinus) adustipennis Erwin, 1969

Brachinus (Neobrachinus) aeger Chaudoir, 1876

Brachinus (Neobrachinus) alexiguus Erwin, 1970

Brachinus (Neobrachinus) alternans Dejean, 1825

Brachinus (Neobrachinus) americanus (LeConte, 1844)

Brachinus (Neobrachinus) azureipennis Chaudoir, 1876

Brachinus (Neobrachinus) capnicus Erwin, 1970

Brachinus (Neobrachinus) cibolensis Erwin, 1970

Brachinus (Neobrachinus) conformis Dejean, 1831

Brachinus (Neobrachinus) cordicollis Dejean, 1826

Brachinus (Neobrachinus) costipennis Motschulsky, 1859

Brachinus (Neobrachinus) cyanipennis Say, 1823

Brachinus (Neobrachinus) cyanochroaticus Erwin, 1969

Brachinus (Neobrachinus) elongatulus Chaudoir, 1876

Brachinus (Neobrachinus) explosus Erwin, 1970

Brachinus (Neobrachinus) favicollis Erwin, 1965

Brachinus (Neobrachinus) fulminatus Erwin, 1969

Brachinus (Neobrachinus) fumans (Fabricius, 1781)

Brachinus (Neobrachinus) gebhardis Erwin, 1965

Brachinus (Neobrachinus) geniculatus Dejean, 1831

Brachinus (Neobrachinus) hirsutus Bates, 1884

Brachinus (Neobrachinus) ichabodopsis Erwin, 1970

Brachinus (Neobrachinus) imperialensis Erwin, 1965

Brachinus (Neobrachinus) imporcitis Erwin, 1970

Brachinus (Neobrachinus) janthinipennis (Dejean, 1831)

Brachinus (Neobrachinus) javalinopsis Erwin, 1970

Brachinus (Neobrachinus) kansanus LeConte, 1863

Brachinus (Neobrachinus) kavanaughi Erwin, 1969

Brachinus (Neobrachinus) lateralis Dejean, 1831

Brachinus (Neobrachinus) medius Harris, 1828

Brachinus (Neobrachinus) mexicanus Dejean, 1831

Brachinus (Neobrachinus) microamericanus Erwin, 1969

Brachinus (Neobrachinus) mobilis Erwin, 1970

Brachinus (Neobrachinus) neglectus LeConte, 1844

Brachinus (Neobrachinus) ovipennis LeConte, 1863

Brachinus (Neobrachinus) oxygonus Chaudoir, 1843

Brachinus (Neobrachinus) pallidus Erwin, 1965

Brachinus (Neobrachinus) patruelis LeConte, 1844

Brachinus (Neobrachinus) perplexus Dejean, 1831

Brachinus (Neobrachinus) phaeocerus Chaudoir, 1868

Brachinus (Neobrachinus) puberulus Chaudoir, 1868

Brachinus (Neobrachinus) quadripennis Dejean, 1825

Brachinus (Neobrachinus) rugipennis Chaudoir, 1868

Brachinus (Neobrachinus) sublaevis Chaudoir, 1868

Brachinus (Neobrachinus) tenuicollis LeConte, 1844

Brachinus (Neobrachinus) texanus Chaudoir, 1868

Brachinus (Neobrachinus) velutinus Erwin, 1965

Brachinus (Neobrachinus) viridipennis Dejean, 1831

Brachinus (Neobrachinus) vulcanoides Erwin, 1969


Morion aridus Allen, 1969

Morion monilicornis (Latreille, 1805)


Loxandrus accelerans Casey, 1918

Loxandrus agilis (Dejean, 1828)

Loxandrus algidus Allen, 1972

Loxandrus brevicollis (LeConte, 1846)

Loxandrus celer (Dejean, 1828)

Loxandrus cervicalis Casey, 1918

Loxandrus cincinnati Casey, 1924

Loxandrus circulus Allen, 1972

Loxandrus collucens Casey, 1918

Loxandrus crenatus LeConte, 1853

Loxandrus duryi Wright, 1939

Loxandrus erraticus (Dejean, 1828)

Loxandrus extendus Allen, 1972

Loxandrus floridanus LeConte, 1878

Loxandrus gibbus Allen, 1972

Loxandrus icarus Will & Liebherr, 1998

Loxandrus infimus Bates, 1882

Loxandrus lucens Chaudoir, 1868

Loxandrus micans Chaudoir, 1868

Loxandrus minor (Chaudoir, 1843)

Loxandrus nitidulus (LeConte, 1846)

Loxandrus pactinullus Allen, 1972

Loxandrus parallelus Casey, 1918

Loxandrus parvulus Chaudoir, 1868

Loxandrus piceolus Chaudoir, 1868

Loxandrus piciventris (LeConte, 1846)

Loxandrus pravitubus Allen, 1972

Loxandrus proximus Chaudoir, 1868

Loxandrus pusillus LeConte, 1853

Loxandrus rectangulus LeConte, 1878

Loxandrus rectus (Say, 1823)

Loxandrus robustus Allen, 1972

Loxandrus rossi Allen, 1972

Loxandrus saccisecundaris Allen, 1972

Loxandrus saphyrinus (Chaudoir, 1843)

Loxandrus sculptilis Bates, 1884

Loxandrus spinilunatus Allen, 1972

Loxandrus straneoi Will & Liebherr, 1998

Loxandrus taeniatus LeConte, 1853

Loxandrus uniformis Allen, 1972

Loxandrus unilobus Allen, 1972

Loxandrus velocipes Casey, 1918

Loxandrus velox (Dejean, 1828)

Loxandrus vulneratus Casey, 1918

Stolonis intercepta Chaudoir, 1874


Abaris (Abaridius) splendidula (LeConte, 1863)

Hybothecus flohri (Bates, 1882)

Poecilus (Poecilus) chalcites (Say, 1823)

Poecilus (Poecilus) coloradensis (Csiki, 1930)

Poecilus (Poecilus) corvus (LeConte, 1873)

Poecilus (Poecilus) cursitor LeConte, 1853

Poecilus (Poecilus) cyanicolor Chaudoir, 1876

Poecilus (Poecilus) diplophryus Chaudoir, 1876

Poecilus (Poecilus) laetulus (LeConte, 1863)

Poecilus (Poecilus) lucublandus (Say, 1823)

Poecilus (Poecilus) mexicanus Chaudoir, 1876

Poecilus (Poecilus) occidentalis (Dejean, 1828)

Poecilus (Poecilus) scitulus LeConte, 1846

Poecilus (Poecilus) texanus (LeConte, 1863)

Poecilus (Derus) nearcticus (Lindroth, 1966)

Lophoglossus gravis LeConte, 1873

Lophoglossus haldemanni (LeConte, 1846)

Lophoglossus scrutator (LeConte, 1846)

Lophoglossus substrenuus (Csiki, 1930)

Lophoglossus tartaricus (Say, 1823)

Lophoglossus vernix Casey, 1913

Piesmus submarginatus (Say, 1823)

Gastrellarius blanchardi (Horn, 1891)

Gastrellarius honestus (Say, 1823)

Gastrellarius unicarum (Darlington, 1932)

Stomis (Neostomis) termitiformis (Van Dyke, 1926)

Stomis (Stomis) pumicatus (Panzer, 1795)

Stereocerus haematopus (Dejean, 1831)

Stereocerus rubripes (Motschulsky, 1860)

Myas (Trigonognatha) coracinus (Say, 1823)

Myas (Trigonognatha) cyanescens Dejean, 1828

Pterostichus (Argutor) commutabilis (Motschulsky, 1866)

Pterostichus (Argutor) praetermissus (Chaudoir, 1868)

Pterostichus (Argutor) vernalis (Panzer, 1795)

Pterostichus (Phonias) corrusculus LeConte, 1873

Pterostichus (Phonias) femoralis (Kirby, 1837)

Pterostichus (Phonias) patruelis (Dejean, 1831)

Pterostichus (Phonias) strenuus (Panzer, 1796)

Pterostichus (Bothriopterus) adstrictus Eschscholtz, 1823

Pterostichus (Bothriopterus) lustrans LeConte, 1851

Pterostichus (Bothriopterus) mutus (Say, 1823)

Pterostichus (Bothriopterus) oregonus LeConte, 1861

Pterostichus (Bothriopterus) pensylvanicus LeConte, 1873

Pterostichus (Bothriopterus) trinarius (Casey, 1918)

Pterostichus (Melanius) castor Goulet & Bousquet, 1983

Pterostichus (Melanius) corvinus (Dejean, 1828)

Pterostichus (Melanius) ebeninus (Dejean, 1828)

Pterostichus (Pseudomaseus) luctuosus (Dejean, 1828)

Pterostichus (Pseudomaseus) tenuis (Casey, 1924)

Pterostichus (Feronina) barri Bousquet, 2006

Pterostichus (Feronina) palmi Schaeffer, 1910

Pterostichus (Paraferonia) lubricus LeConte, 1853

Pterostichus (Pseudoferonina) amadeus Bousquet, nomen novum

Pterostichus (Pseudoferonina) bousqueti Bergdahl, 2011

Pterostichus (Pseudoferonina) campbelli Bousquet, 1985

Pterostichus (Pseudoferonina) humidulus (Van Dyke, 1943)

Pterostichus (Pseudoferonina) lanei Van Dyke, 1926

Pterostichus (Pseudoferonina) lolo Bergdahl, 2011

Pterostichus (Pseudoferonina) shulli (Hatch, 1949)

Pterostichus (Pseudoferonina) smetanai Bousquet, 1985

Pterostichus (Pseudoferonina) spathifer Bousquet, 1992

Pterostichus (Gastrosticta) enodis Bousquet, 1992

Pterostichus (Gastrosticta) mutoides Bousquet, 1992

Pterostichus (Gastrosticta) obesulus LeConte, 1873

Pterostichus (Gastrosticta) ophryoderus (Chaudoir, 1878)

Pterostichus (Gastrosticta) punctiventris (Chaudoir, 1878)

Pterostichus (Gastrosticta) putus Casey, 1913

Pterostichus (Gastrosticta) sayanus Csiki, 1930

Pterostichus (Gastrosticta) subacutus (Casey, 1918)

Pterostichus (Gastrosticta) tumescens LeConte, 1863

Pterostichus (Gastrosticta) ventralis (Say, 1823)

Pterostichus (Morphnosoma) melanarius melanarius (Illiger, 1798)

Pterostichus (Euferonia) coracinus (Newman, 1838)

Pterostichus (Euferonia) ingens (Casey, 1918)

Pterostichus (Euferonia) lachrymosus (Newman, 1838)

Pterostichus (Euferonia) novus Straneo, 1944

Pterostichus (Euferonia) relictus (Newman, 1838)

Pterostichus (Euferonia) stygicus (Say, 1823)

Pterostichus (Lenapterus) agonus Horn, 1880

Pterostichus (Lenapterus) costatus (Ménétriés, 1851)

Pterostichus (Lenapterus) punctatissimus (Randall, 1838)

Pterostichus (Lenapterus) vermiculosus (Ménétriés, 1851)

Pterostichus (Metallophilus) sublaevis (Sahlberg, 1880)

Pterostichus (Abacidus) atratus (Newman, 1838)

Pterostichus (Abacidus) fallax (Dejean, 1828)

Pterostichus (Abacidus) hamiltoni Horn, 1880

Pterostichus (Abacidus) permundus (Say, 1830)

Pterostichus (Abacidus) sculptus LeConte, 1853

Pterostichus (Orsonjohnsonus) johnsoni Ulke, 1889

Pterostichus (Lamenius) caudicalis (Say, 1823)

Pterostichus (Eosteropus) circulosus Lindroth, 1966

Pterostichus (Eosteropus) moestus (Say, 1823)

Pterostichus (Eosteropus) superciliosus (Say, 1823)

Pterostichus (Monoferonia) carolinus carolinus Darlington, 1932

Pterostichus (Monoferonia) carolinus fumorum Darlington, 1932

Pterostichus (Monoferonia) diligendus (Chaudoir, 1868)

Pterostichus (Monoferonia) mancus (LeConte, 1853)

Pterostichus (Monoferonia) primus Darlington, 1932

Pterostichus (Cylindrocharis) acutipes acutipes Barr, 1971

Pterostichus (Cylindrocharis) acutipes kentuckensis Barr, 1971

Pterostichus (Cylindrocharis) hypogeus Barr, 1971

Pterostichus (Cylindrocharis) rostratus (Newman, 1838)

Pterostichus (Leptoferonia) angustus (Dejean, 1828)

Pterostichus (Leptoferonia) beyeri Van Dyke, 1926

Pterostichus (Leptoferonia) blodgettensis Will, 2007

Pterostichus (Leptoferonia) caligans Horn, 1891

Pterostichus (Leptoferonia) cochlearis Hacker, 1968

Pterostichus (Leptoferonia) deino Will, 2007

Pterostichus (Leptoferonia) enyo Will, 2007

Pterostichus (Leptoferonia) falli Van Dyke, 1926

Pterostichus (Leptoferonia) fenyesi fenderi Hacker, 1968

Pterostichus (Leptoferonia) fenyesi fenyesi Csiki, 1930

Pterostichus (Leptoferonia) fuchsi Schaeffer, 1910

Pterostichus (Leptoferonia) hatchi Hacker, 1968

Pterostichus (Leptoferonia) humilis Casey, 1913

Pterostichus (Leptoferonia) idahoae Csiki, 1930

Pterostichus (Leptoferonia) inanis Horn, 1891

Pterostichus (Leptoferonia) infernalis Hatch, 1936

Pterostichus (Leptoferonia) inopinus (Casey, 1918)

Pterostichus (Leptoferonia) lobatus Hacker, 1968

Pterostichus (Leptoferonia) marinensis Hacker, 1968

Pterostichus (Leptoferonia) mattolensis Hacker, 1968

Pterostichus (Leptoferonia) pemphredo Will, 2007

Pterostichus (Leptoferonia) pumilus pumilus Casey, 1913

Pterostichus (Leptoferonia) pumilus willamettensis Hacker, 1968

Pterostichus (Leptoferonia) rothi (Hatch, 1951)

Pterostichus (Leptoferonia) sphodrinus LeConte, 1863

Pterostichus (Leptoferonia) stapedius Hacker, 1968

Pterostichus (Leptoferonia) trinitensis Hacker, 1968

Pterostichus (Leptoferonia) yosemitensis Hacker, 1968

Pterostichus (Anilloferonia) lanei (Hatch, 1935)

Pterostichus (Anilloferonia) malkini (Hatch, 1953)

Pterostichus (Anilloferonia) testaceus (Van Dyke, 1926)

Pterostichus (Hypherpes) adoxus (Say, 1823)

Pterostichus (Hypherpes) algidus LeConte, 1853

Pterostichus (Hypherpes) amethystinus Mannerheim, 1843

Pterostichus (Hypherpes) annosus Casey, 1913

Pterostichus (Hypherpes) arcanus Casey, 1913

Pterostichus (Hypherpes) baldwini (Casey, 1924)

Pterostichus (Hypherpes) barbarinus Casey, 1913

Pterostichus (Hypherpes) brachylobus Kavanaugh & LaBonte, 2006

Pterostichus (Hypherpes) californicus (Dejean, 1828)

Pterostichus (Hypherpes) canallatus Casey, 1913

Pterostichus (Hypherpes) castaneus (Dejean, 1828)

Pterostichus (Hypherpes) castanipes (Ménétriés, 1843)

Pterostichus (Hypherpes) congestus (Ménétriés, 1843)

Pterostichus (Hypherpes) craterensis (Hatch, 1949)

Pterostichus (Hypherpes) crenicollis LeConte, 1873

Pterostichus (Hypherpes) ecarinatus Hatch, 1936

Pterostichus (Hypherpes) esuriens Casey, 1913

Pterostichus (Hypherpes) gliscans Casey, 1913

Pterostichus (Hypherpes) gracilior LeConte, 1873

Pterostichus (Hypherpes) herculaneus Mannerheim, 1843

Pterostichus (Hypherpes) hornii LeConte, 1873

Pterostichus (Hypherpes) illustris LeConte, 1851

Pterostichus (Hypherpes) inermis Fall, 1901

Pterostichus (Hypherpes) isabellae LeConte, 1851

Pterostichus (Hypherpes) jacobinus Casey, 1913

Pterostichus (Hypherpes) laborans Casey, 1913

Pterostichus (Hypherpes) lacertus Casey, 1913

Pterostichus (Hypherpes) lama (Ménétriés, 1843)

Pterostichus (Hypherpes) lassulus (Casey, 1920)

Pterostichus (Hypherpes) lattini LaBonte, 2006

Pterostichus (Hypherpes) luscus (Casey, 1918)

Pterostichus (Hypherpes) menetriesii LeConte, 1873

Pterostichus (Hypherpes) mercedianus (Casey, 1918)

Pterostichus (Hypherpes) miscellus Casey, 1913

Pterostichus (Hypherpes) morionides (Chaudoir, 1868)

Pterostichus (Hypherpes) neobrunneus Lindroth, 1966

Pterostichus (Hypherpes) nigrocaeruleus Van Dyke, 1926

Pterostichus (Hypherpes) obsidianus Casey, 1913

Pterostichus (Hypherpes) occultus Casey, 1913

Pterostichus (Hypherpes) ordinarius Casey, 1913

Pterostichus (Hypherpes) ovalipennis Casey, 1913

Pterostichus (Hypherpes) panticulatus Casey, 1913

Pterostichus (Hypherpes) pergracilis (Casey, 1920)

Pterostichus (Hypherpes) planctus LeConte, 1853

Pterostichus (Hypherpes) protensiformis (Casey, 1924)

Pterostichus (Hypherpes) protractus LeConte, 1860

Pterostichus (Hypherpes) restrictus (Casey, 1918)

Pterostichus (Hypherpes) scutellaris LeConte, 1873

Pterostichus (Hypherpes) serripes (LeConte, 1875)

Pterostichus (Hypherpes) setosus Hatch, 1951

Pterostichus (Hypherpes) sierranus Casey, 1913

Pterostichus (Hypherpes) sponsor Casey, 1913

Pterostichus (Hypherpes) spraguei LeConte, 1873

Pterostichus (Hypherpes) suffusus Casey, 1913

Pterostichus (Hypherpes) tarsalis LeConte, 1873

Pterostichus (Hypherpes) tristis (Dejean, 1828)

Pterostichus (Hypherpes) tuberculofemoratus Hatch, 1936

Pterostichus (Hypherpes) vandykei Schaeffer, 1910

Pterostichus (Hypherpes) vicinus Mannerheim, 1843

Pterostichus (Hypherpes) ybousqueti Berlov, 1999

Pterostichus (Cryobius) arcticola (Chaudoir, 1868)

Pterostichus (Cryobius) auriga Ball, 1962

Pterostichus (Cryobius) barryorum Ball, 1962

Pterostichus (Cryobius) brevicornis brevicornis (Kirby, 1837)

Pterostichus (Cryobius) bryanti biocryus Ball, 1962

Pterostichus (Cryobius) bryanti bryanti (Van Dyke, 1951)

Pterostichus (Cryobius) bryanti bryantoides Ball, 1962

Pterostichus (Cryobius) bryanti cacumenis Ball, 1966

Pterostichus (Cryobius) bryanti stantonensis Ball, 1966

Pterostichus (Cryobius) bryanti tiliaceoradix Ball, 1962

Pterostichus (Cryobius) caribou Ball, 1962

Pterostichus (Cryobius) chipewyan Ball, 1962

Pterostichus (Cryobius) empetricola (Dejean, 1828)

Pterostichus (Cryobius) gerstlensis Ball, 1962

Pterostichus (Cryobius) hudsonicus LeConte, 1863

Pterostichus (Cryobius) kotzebuei Ball, 1962

Pterostichus (Cryobius) mandibularoides Ball, 1966

Pterostichus (Cryobius) nivalis (Sahlberg, 1844)

Pterostichus (Cryobius) parasimilis Ball, 1962

Pterostichus (Cryobius) pinguedineus (Eschscholtz, 1823)

Pterostichus (Cryobius) planus (Sahlberg, 1885)

Pterostichus (Cryobius) riparius (Dejean, 1828)

Pterostichus (Cryobius) similis Mannerheim, 1852

Pterostichus (Cryobius) soperi Ball, 1966

Pterostichus (Cryobius) surgens LeConte, 1878

Pterostichus (Cryobius) tareumiut Ball, 1962

Pterostichus (Cryobius) ventricosus ventricosus (Eschscholtz, 1823)

Pterostichus (Cryobius) woodi Ball & Currie, 1997

Cyclotrachelus (Cyclotrachelus) alabamensis (Casey, 1920)

Cyclotrachelus (Cyclotrachelus) approximatus (LeConte, 1846)

Cyclotrachelus (Cyclotrachelus) brevoorti (LeConte, 1846)

Cyclotrachelus (Cyclotrachelus) dejeanellus (Csiki, 1930)

Cyclotrachelus (Cyclotrachelus) faber (Germar, 1824)

Cyclotrachelus (Cyclotrachelus) freitagi Bousquet, 1993

Cyclotrachelus (Cyclotrachelus) fucatus (Freitag, 1969)

Cyclotrachelus (Cyclotrachelus) hernandensis (Van Dyke, 1943)

Cyclotrachelus (Cyclotrachelus) iuvenis (Freitag, 1969)

Cyclotrachelus (Cyclotrachelus) laevipennis (LeConte, 1846)

Cyclotrachelus (Cyclotrachelus) levifaber (Freitag, 1969)

Cyclotrachelus (Cyclotrachelus) macrovulum (Freitag, 1969)

Cyclotrachelus (Cyclotrachelus) ovulum (Chaudoir, 1868)

Cyclotrachelus (Cyclotrachelus) parafaber (Freitag, 1969)

Cyclotrachelus (Cyclotrachelus) spoliatus (Newman, 1838)

Cyclotrachelus (Cyclotrachelus) texensis (Freitag, 1969)

Cyclotrachelus (Cyclotrachelus) unicolor (Say, 1823)

Cyclotrachelus (Cyclotrachelus) vinctus (LeConte, 1853)

Cyclotrachelus (Evarthrus) alabamae (Van Dyke, 1926)

Cyclotrachelus (Evarthrus) alternans (Casey, 1920)

Cyclotrachelus (Evarthrus) blatchleyi (Casey, 1918)

Cyclotrachelus (Evarthrus) constrictus (Say, 1823)

Cyclotrachelus (Evarthrus) convivus (LeConte, 1853)

Cyclotrachelus (Evarthrus) deceptus (Casey, 1918)

Cyclotrachelus (Evarthrus) engelmani (LeConte, 1853)

Cyclotrachelus (Evarthrus) floridensis (Freitag, 1969)

Cyclotrachelus (Evarthrus) furtivus (LeConte, 1853)

Cyclotrachelus (Evarthrus) gigas (Casey, 1918)

Cyclotrachelus (Evarthrus) gravesi (Freitag, 1969)

Cyclotrachelus (Evarthrus) gravidus (Haldeman, 1853)

Cyclotrachelus (Evarthrus) heros (Say, 1823)

Cyclotrachelus (Evarthrus) hypherpiformis (Freitag, 1969)

Cyclotrachelus (Evarthrus) incisus (LeConte, 1846)

Cyclotrachelus (Evarthrus) iowensis (Freitag, 1969)

Cyclotrachelus (Evarthrus) lodingi (Van Dyke, 1926)

Cyclotrachelus (Evarthrus) nonnitens (LeConte, 1873)

Cyclotrachelus (Evarthrus) parasodalis (Freitag, 1969)

Cyclotrachelus (Evarthrus) sallei (LeConte, 1873)

Cyclotrachelus (Evarthrus) seximpressus (LeConte, 1846)

Cyclotrachelus (Evarthrus) sigillatus (Say, 1823)

Cyclotrachelus (Evarthrus) sinus (Freitag, 1969)

Cyclotrachelus (Evarthrus) sodalis colossus (LeConte, 1846)

Cyclotrachelus (Evarthrus) sodalis sodalis (LeConte, 1846)

Cyclotrachelus (Evarthrus) substriatus (LeConte, 1846)

Cyclotrachelus (Evarthrus) torvus (LeConte, 1863)

Cyclotrachelus (Evarthrus) whitcombi (Freitag, 1969)

Abax (Abax) parallelepipedus (Piller & Mitterpacher, 1783)


Amara (Curtonotus) alpina (Paykull, l790)

Amara (Curtonotus) aulica (Panzer, 1796)

Amara (Curtonotus) blanchardi Hayward, 1908

Amara (Curtonotus) bokori Csiki, 1929

Amara (Curtonotus) carinata (LeConte, 1847)

Amara (Curtonotus) daurica (Motschulsky, 1844)

Amara (Curtonotus) deparca (Say, 1830)

Amara (Curtonotus) hyperborea Dejean, 1831

Amara (Curtonotus) jacobina LeConte, 1855

Amara (Curtonotus) kurnakowi Hieke, 1994

Amara (Curtonotus) lacustris LeConte, 1855

Amara (Curtonotus) pennsylvanica Hayward, 1908

Amara (Curtonotus) pterostichina Hayward, 1908

Amara (Curtonotus) thoracica Hayward, 1908

Amara (Curtonotus) torrida (Panzer, 1796)

Amara (Bradytus) apricaria (Paykull, 1790)

Amara (Bradytus) avida (Say, 1823)

Amara (Bradytus) browni Lindroth, 1968

Amara (Bradytus) exarata Dejean, 1828

Amara (Bradytus) fulva (Müller, 1776)

Amara (Bradytus) glacialis (Mannerheim, 1853)

Amara (Bradytus) insignis Dejean, 1831

Amara (Bradytus) insularis Horn, 1875

Amara (Bradytus) latior (Kirby, 1837)

Amara (Bradytus) lindrothi Hieke, 1990

Amara (Bradytus) neomexicana (Casey, 1924)

Amara (Bradytus) schwarzi Hayward, 1908

Amara (Neopercosia) fortis LeConte, 1880

Amara (Percosia) obesa (Say, 1823)

Amara (Xenocelia) apachensis Casey, 1884

Amara (Xenocelia) bradytonota Hieke, 2001

Amara (Xenocelia) chalcea Dejean, 1828

Amara (Xenocelia) discors Kirby, 1837

Amara (Xenocelia) gibba (LeConte, 1847)

Amara (Xenocelia) harpalonota Hieke, 2001

Amara (Xenocelia) hicksi Lindroth, 1968

Amara (Xenocelia) lugubris (Casey, 1918)

Amara (Xenocelia) merula (Casey, 1918)

Amara (Xenocelia) rectangula ciudadensis (Bates, 1891)

Amara (Xenocelia) rectangula rectangula LeConte, 1855

Amara (Xenocelia) spuria Lindroth, 1968

Amara (Reductocelia) colvillensis Lindroth, 1968

Amara (Celia) bifrons (Gyllenhal, 1810)

Amara (Celia) brunnea (Gyllenhal, 1810)

Amara (Celia) californica californica Dejean, 1828

Amara (Celia) exlineae Minsk & Hatch, 1939

Amara (Celia) harpalina LeConte, 1855

Amara (Celia) idahoana (Casey, 1924)

Amara (Celia) musculis (Say, 1823)

Amara (Celia) pseudobrunnea Lindroth, 1968

Amara (Celia) rubrica Haldeman, 1843

Amara (Celia) sinuosa (Casey, 1918)

Amara (Celia) texana (Putzeys, 1866)

Amara (Celia) volatilis (Casey, 1918)

Amara (Amarocelia) ellipsis (Casey, 1918)

Amara (Amarocelia) erratica (Duftschmid, 1812)

Amara (Amarocelia) farcta LeConte, 1855

Amara (Amarocelia) interstitialis Dejean, 1828

Amara (Amarocelia) laevipennis Kirby, 1837

Amara (Amarocelia) lugens Zimmermann, 1832

Amara (Amarocelia) nexa (Casey, 1918)

Amara (Amarocelia) patruelis Dejean, 1831

Amara (Amarocelia) rugulifera Hieke, 2002

Amara (Amarocelia) sodalicia Casey, 1924

Amara (Amarocelia) tenebrionella (Bates, 1882)

Amara (Amarocelia) transberingiensis Hieke, 2002

Amara (Amara) aenea (DeGeer, 1774)

Amara (Amara) aeneopolita Casey, 1918

Amara (Amara) anthobia Villa & Villa, 1833

Amara (Amara) aurata Dejean, 1828

Amara (Amara) basillaris (Say, 1823)

Amara (Amara) coelebs Hayward, 1908

Amara (Amara) communis (Panzer, 1797)

Amara (Amara) conflata LeConte, 1855

Amara (Amara) confusa LeConte, 1847

Amara (Amara) convexa LeConte, 1847

Amara (Amara) crassispina LeConte, 1855

Amara (Amara) cupreolata Putzeys, 1866

Amara (Amara) emancipata Lindroth, 1968

Amara (Amara) eurynota (Panzer, 1796)

Amara (Amara) externefoveata Hieke, 2002

Amara (Amara) familiaris (Duftschmid, 1812)

Amara (Amara) haywardi Csiki, 1929

Amara (Amara) impuncticollis (Say, 1823)

Amara (Amara) littoralis Dejean, 1828

Amara (Amara) lunicollis Schiødte, 1837

Amara (Amara) neoscotica Casey, 1924

Amara (Amara) occidentalis Hieke, 2002

Amara (Amara) otiosa Casey, 1918

Amara (Amara) ovata (Fabricius, 1792)

Amara (Amara) pomona Casey, 1918

Amara (Amara) sanjuanensis Hatch, 1949

Amara (Amara) sera Say, 1830

Amara (Amara) tenax Casey, 1918

Amara (Amara) turbata Casey, 1918

Amara (Paracelia) quenseli quenseli (Schönherr, 1806)

Amara (Zezea) angustata (Say, 1823)

Amara (Zezea) angustatoides Hieke, 2000

Amara (Zezea) belfragei Horn, 1892

Amara (Zezea) flebilis (Casey, 1918)

Amara (Zezea) inexspectata Hieke, 1990

Amara (Zezea) kavanaughi Hieke, 1990

Amara (Zezea) longula LeConte, 1855

Amara (Zezea) pallipes Kirby, 1837

Amara (Zezea) scitula Zimmermann, 1832


Dercylinus impressus (LeConte, 1853)

Evolenes exarata (Dejean, 1831)

Anatrichis minuta (Dejean, 1831)

Anatrichis oblonga Horn, 1891

Oodinus alutaceus (Bates, 1882)

Oodinus pseudopiceus Bousquet, 1996

Lachnocrepis parallela (Say, 1830)

Oodes amaroides Dejean, 1831

Oodes americanus Dejean, 1826

Oodes brevis Lindroth, 1957

Oodes fluvialis LeConte, 1863

Stenocrepis (Stenocrepis) insulana (Jacquelin du Val, 1857)

Stenocrepis (Stenous) cuprea (Chaudoir, 1843)

Stenocrepis (Stenous) duodecimstriata (Chevrolat, 1836)

Stenocrepis (Stenous) elegans (LeConte, 1851)

Stenocrepis (Stenous) mexicana (Chevrolat, 1835)

Stenocrepis (Stenous) tibialis (Chevrolat, 1834)


Panagaeus (Hologaeus) cruciger Say, 1823

Panagaeus (Hologaeus) fasciatus Say, 1823

Panagaeus (Hologaeus) sallei Chaudoir, 1862

Micrixys distincta (Haldeman, 1852)


Chlaenius (Pseudanomoglossus) maxillosus Horn, 1876

Chlaenius (Eurydactylus) pimalicus Casey, 1914

Chlaenius (Eurydactylus) tomentosus (Say, 1823)

Chlaenius (Anomoglossus) amoenus Dejean, 1831

Chlaenius (Anomoglossus) emarginatus Say, 1823

Chlaenius (Anomoglossus) pusillus Say, 1823

Chlaenius (Chlaenius) aestivus Say, 1823

Chlaenius (Chlaenius) augustus Newman, 1838

Chlaenius (Chlaenius) azurescens Chaudoir, 1876

Chlaenius (Chlaenius) chaudoiri Horn, 1876

Chlaenius (Chlaenius) cumatilis LeConte, 1851

Chlaenius (Chlaenius) erythropus Germar, 1824

Chlaenius (Chlaenius) fuscicornis Dejean, 1831

Chlaenius (Chlaenius) laticollis Say, 1823

Chlaenius (Chlaenius) orbus Horn, 1871

Chlaenius (Chlaenius) patruelis LeConte, 1844

Chlaenius (Chlaenius) platyderus Chaudoir, 1856

Chlaenius (Chlaenius) sericeus (Forster, 1771)

Chlaenius (Chlaenius) sparsus LeConte, 1863

Chlaenius (Chlaenius) viduus Horn, 1871

Chlaenius (Lithochlaenius) cordicollis Kirby, 1837

Chlaenius (Lithochlaenius) leucoscelis monachus LeConte, 1851

Chlaenius (Lithochlaenius) leucoscelis sanantonialis Casey, 1914

Chlaenius (Lithochlaenius) leucoscelis sonomae Casey, 1920

Chlaenius (Lithochlaenius) prasinus Dejean, 1826

Chlaenius (Lithochlaenius) purpureus Chaudoir, 1876

Chlaenius (Lithochlaenius) solitarius Say, 1823

Chlaenius (Chlaeniellus) brevilabris LeConte, 1847

Chlaenius (Chlaeniellus) circumcinctus Say, 1830

Chlaenius (Chlaeniellus) flaccidus Horn, 1876

Chlaenius (Chlaeniellus) floridanus Horn, 1876

Chlaenius (Chlaeniellus) glaucus LeConte, 1856

Chlaenius (Chlaeniellus) impunctifrons Say, 1823

Chlaenius (Chlaeniellus) nebraskensis LeConte, 1856

Chlaenius (Chlaeniellus) nemoralis Say, 1823

Chlaenius (Chlaeniellus) obsoletus LeConte, 1851

Chlaenius (Chlaeniellus) oxygonus Chaudoir, 1843

Chlaenius (Chlaeniellus) pennsylvanicus blanditus Casey, 1920

Chlaenius (Chlaeniellus) pennsylvanicus pennsylvanicus Say, 1823

Chlaenius (Chlaeniellus) pertinax Casey, 1920

Chlaenius (Chlaeniellus) simillimus Chaudoir, 1856

Chlaenius (Chlaeniellus) texanus Horn, 1876

Chlaenius (Chlaeniellus) tricolor tricolor Dejean, 1826

Chlaenius (Chlaeniellus) tricolor vigilans Say, 1830

Chlaenius (Chlaeniellus) vafer LeConte, 1852

Chlaenius (Chlaeniellus) variabilipes Eschscholtz, 1833

Chlaenius (Callistometus) ruficauda Chaudoir, 1856

Chlaenius (Brachylobus) caurinus (Horn, 1885)

Chlaenius (Brachylobus) lithophilus Say, 1823

Chlaenius (Agostenus) alternatus Horn, 1871

Chlaenius (Agostenus) caeruleicollis Chaudoir, 1876

Chlaenius (Agostenus) harpalinus Eschscholtz, 1833

Chlaenius (Agostenus) interruptus Horn, 1876

Chlaenius (Agostenus) niger Randall, 1838

Chlaenius (Randallius) purpuricollis Randall, 1838


Diplocheila (Isorembus) assimilis (LeConte, 1844)

Diplocheila (Isorembus) crossi Will, 1998

Diplocheila (Isorembus) impressicollis (Dejean, 1831)

Diplocheila (Isorembus) major major (LeConte, 1847)

Diplocheila (Isorembus) major melissisa Ball, 1959

Diplocheila (Isorembus) nupera Casey, 1897

Diplocheila (Isorembus) obtusa (LeConte, 1847)

Diplocheila (Isorembus) oregona (Hatch, 1951)

Diplocheila (Isorembus) striatopunctata (LeConte, 1844)

Diplocheila (Isorembus) undulata Carr, 1920

Dicaelus (Paradicaelus) ambiguus LaFerté-Sénectère, 1841

Dicaelus (Paradicaelus) dilatatus dilatatus Say, 1823

Dicaelus (Paradicaelus) dilatatus sinuatus Ball, 1959

Dicaelus (Paradicaelus) elongatus Bonelli, 1813

Dicaelus (Paradicaelus) furvus carinatus Dejean, 1831

Dicaelus (Paradicaelus) furvus furvus Dejean, 1826

Dicaelus (Paradicaelus) politus Dejean, 1826

Dicaelus (Paradicaelus) sculptilis intricatus LeConte, 1873

Dicaelus (Paradicaelus) sculptilis sculptilis Say, 1823

Dicaelus (Paradicaelus) sculptilis upioides Ball, 1959

Dicaelus (Paradicaelus) teter Bonelli, 1813

Dicaelus (Dicaelus) alternans Dejean, 1826

Dicaelus (Dicaelus) costatus LeConte, 1853

Dicaelus (Dicaelus) crenatus LeConte, 1853

Dicaelus (Dicaelus) purpuratus purpuratus Bonelli, 1813

Dicaelus (Dicaelus) purpuratus splendidus Say, 1823

Dicaelus (Dicaelus) quadratus LeConte, 1847

Dicaelus (Dicaelus) subtropicus Casey, 1913

Dicaelus (Liodicaelus) chermocki Ball, 1959

Dicaelus (Liodicaelus) laevipennis laevipennis LeConte, 1847

Dicaelus (Liodicaelus) suffusus (Casey, 1913)

Badister (Badister) elegans LeConte, 1880

Badister (Badister) ferrugineus Dejean, 1831

Badister (Badister) flavipes flavipes LeConte, 1853

Badister (Badister) maculatus LeConte, 1853

Badister (Badister) neopulchellus Lindroth, 1954

Badister (Badister) notatus Haldeman, 1843

Badister (Badister) obtusus LeConte, 1878

Badister (Badister) pulchellus LeConte, 1847

Badister (Baudia) grandiceps Casey, 1920

Badister (Baudia) micans LeConte, 1844

Badister (Baudia) parviceps Ball, 1959

Badister (Baudia) reflexus LeConte, 1880

Badister (Baudia) submarinus Motschulsky, 1859

Badister (Baudia) transversus Casey, 1920


Notiobia (Anisotarsus) brevicollis (Chaudoir, 1837)

Notiobia (Anisotarsus) cephala (Casey, 1914)

Notiobia (Anisotarsus) maculicornis (Chaudoir, 1843)

Notiobia (Anisotarsus) mexicana (Dejean, 1829)

Notiobia (Anisotarsus) nitidipennis (LeConte, 1847)

Notiobia (Anisotarsus) purpurascens (Bates, 1882)

Notiobia (Anisotarsus) sayi (Blatchley, 1910)

Notiobia (Anisotarsus) terminata (Say, 1823)

Xestonotus lugubris (Dejean, 1829)

Anisodactylus (Anisodactylus) agricola (Say, 1823)

Anisodactylus (Anisodactylus) binotatus (Fabricius, 1787)

Anisodactylus (Anisodactylus) californicus Dejean, 1829

Anisodactylus (Anisodactylus) carbonarius (Say, 1823)

Anisodactylus (Anisodactylus) consobrinus LeConte, 1851

Anisodactylus (Anisodactylus) furvus LeConte, 1863

Anisodactylus (Anisodactylus) harrisii LeConte, 1863

Anisodactylus (Anisodactylus) kirbyi Lindroth, 1953

Anisodactylus (Anisodactylus) lodingi Schaeffer, 1911

Anisodactylus (Anisodactylus) melanopus (Haldeman, 1843)

Anisodactylus (Anisodactylus) nigerrimus (Dejean, 1831)

Anisodactylus (Anisodactylus) nigrita Dejean, 1829

Anisodactylus (Anisodactylus) pseudagricola Noonan, 1996

Anisodactylus (Anisodactylus) similis LeConte, 1851

Anisodactylus (Gynandrotarsus) anthracinus (Dejean, 1829)

Anisodactylus (Gynandrotarsus) dulcicollis (LaFerté-Sénectère, 1841)

Anisodactylus (Gynandrotarsus) haplomus Chaudoir, 1868

Anisodactylus (Gynandrotarsus) harpaloides (LaFerté-Sénectère, 1841)

Anisodactylus (Gynandrotarsus) merula (Germar, 1824)

Anisodactylus (Gynandrotarsus) opaculus (LeConte, 1863)

Anisodactylus (Gynandrotarsus) ovularis (Casey, 1914)

Anisodactylus (Gynandrotarsus) rusticus (Say, 1823)

Anisodactylus (Gynandrotarsus) texanus Schaeffer, 1910

Anisodactylus (Anadaptus) alternans (Motschulsky, 1845)

Anisodactylus (Anadaptus) discoideus Dejean, 1831

Anisodactylus (Anadaptus) pitychrous LeConte, 1861

Anisodactylus (Anadaptus) porosus (Motschulsky, 1845)

Anisodactylus (Anadaptus) rudis LeConte, 1863

Anisodactylus (Anadaptus) sanctaecrucis (Fabricius, 1798)

Anisodactylus (Spongopus) verticalis (LeConte, 1847)

Anisodactylus (Aplocentrus) amaroides LeConte, 1851

Anisodactylus (Aplocentrus) caenus (Say, 1823)

Anisodactylus (Pseudaplocentrus) laetus Dejean, 1829

Geopinus incrassatus (Dejean, 1829)

Amphasia (Pseudamphasia) sericea (Harris, 1828)

Amphasia (Amphasia) interstitialis (Say, 1823)

Dicheirus brunneus (Dejean, 1829)

Dicheirus dilatatus angulatus Casey, 1914

Dicheirus dilatatus dilatatus (Dejean, 1829)

Dicheirus obtusus LeConte, 1852

Dicheirus piceus (Ménétriés, 1843)

Dicheirus strenuus (Horn, 1869)

Pelmatellus (Pelmatellus) obtusus Bates, 1882

Pelmatellus (Pelmatellus) stenolophoides parallelus Goulet, 1974

Stenolophus (Stenolophus) anceps LeConte, 1857

Stenolophus (Stenolophus) carbo Bousquet, 1993

Stenolophus (Stenolophus) cincticollis LeConte, 1858

Stenolophus (Stenolophus) dissimilis Dejean, 1829

Stenolophus (Stenolophus) flavipes LeConte, 1858

Stenolophus (Stenolophus) fuliginosus Dejean, 1829

Stenolophus (Stenolophus) fuscatus Dejean, 1829

Stenolophus (Stenolophus) humidus Hamilton, 1893

Stenolophus (Stenolophus) incultus Casey, 1914

Stenolophus (Stenolophus) limbalis LeConte, 1857

Stenolophus (Stenolophus) megacephalus Lindroth, 1968

Stenolophus (Stenolophus) ochropezus (Say, 1823)

Stenolophus (Stenolophus) plebejus Dejean, 1829

Stenolophus (Stenolophus) splendidulus Motschulsky, 1864

Stenolophus (Stenolophus) spretus Dejean, 1831

Stenolophus (Agonoderus) binotatus (Casey, 1914)

Stenolophus (Agonoderus) comma (Fabricius, 1775)

Stenolophus (Agonoderus) infuscatus (Dejean, 1829)

Stenolophus (Agonoderus) lecontei (Chaudoir, 1868)

Stenolophus (Agonoderus) lineola (Fabricius, 1775)

Stenolophus (Agonoderus) maculatus (LeConte, 1869)

Stenolophus (Agonoderus) rugicollis (LeConte, 1859)

Agonoleptus conjunctus (Say, 1823)

Agonoleptus dolosus (Casey, 1914)

Agonoleptus parviceps Casey, 1914

Agonoleptus rotundatus (LeConte, 1863)

Agonoleptus rotundicollis (Haldeman, 1843)

Agonoleptus thoracicus (Casey, 1914)

Agonoleptus unicolor (Dejean, 1829)

Bradycellus (Liocellus) curticollis (Casey, 1924)

Bradycellus (Liocellus) intermedius (Fall, 1905)

Bradycellus (Liocellus) laticollis (Casey, 1924)

Bradycellus (Liocellus) nitidus (Dejean, 1829)

Bradycellus (Liocellus) obtusus (Fall, 1905)

Bradycellus (Liocellus) politus (Fall, 1905)

Bradycellus (Liocellus) tahoensis (Casey, 1924)

Bradycellus (Bradycellus) fenderi Hatch, 1951

Bradycellus (Bradycellus) harpalinus (Audinet-Serville, 1821)

Bradycellus (Catharellus) lecontei Csiki, 1932

Bradycellus (Stenocellus) ardelio (Casey, 1914)

Bradycellus (Stenocellus) aridus (Casey, 1914)

Bradycellus (Stenocellus) californicus (LeConte, 1857)

Bradycellus (Stenocellus) carolinensis (Casey, 1924)

Bradycellus (Stenocellus) congener (LeConte, 1847)

Bradycellus (Stenocellus) decorus (Casey, 1914)

Bradycellus (Stenocellus) discipulus (Casey, 1914)

Bradycellus (Stenocellus) exstans (Casey, 1914)

Bradycellus (Stenocellus) festinans (Casey, 1914)

Bradycellus (Stenocellus) humboldtianus (Casey, 1924)

Bradycellus (Stenocellus) insulsus (Casey, 1914)

Bradycellus (Stenocellus) larvatus (Casey, 1914)

Bradycellus (Stenocellus) lineatus (Casey, 1914)

Bradycellus (Stenocellus) lustrellus (Casey, 1914)

Bradycellus (Stenocellus) montanus (Casey, 1914)

Bradycellus (Stenocellus) nebulosus LeConte, 1853

Bradycellus (Stenocellus) neglectus (LeConte, 1847)

Bradycellus (Stenocellus) nigerrimus Lindroth, 1968

Bradycellus (Stenocellus) nigriceps LeConte, 1869

Bradycellus (Stenocellus) nubifer LeConte, 1858

Bradycellus (Stenocellus) picipes (Casey, 1914)

Bradycellus (Stenocellus) provoensis (Casey, 1914)

Bradycellus (Stenocellus) puncticollis (Casey, 1914)

Bradycellus (Stenocellus) purgatus (Casey, 1914)

Bradycellus (Stenocellus) rivalis LeConte, 1858

Bradycellus (Stenocellus) rupestris (Say, 1823)

Bradycellus (Stenocellus) sejunctus (Casey, 1914)

Bradycellus (Stenocellus) suavis (Casey, 1914)

Bradycellus (Stenocellus) subcordatus Chaudoir, 1868

Bradycellus (Stenocellus) supplex (Casey, 1914)

Bradycellus (Stenocellus) symetricus (Motschulsky, 1850)

Bradycellus (Stenocellus) tantillus (Dejean, 1829)

Bradycellus (Stenocellus) veronianus (Casey, 1924)

Bradycellus (Lipalocellus) nigrinus (Dejean, 1829)

Bradycellus (Lipalocellus) semipubescens Lindroth, 1968

Bradycellus (Triliarthrus) atrimedeus (Say, 1823)

Bradycellus (Triliarthrus) badipennis (Haldeman, 1843)

Bradycellus (Triliarthrus) conformis (Fall, 1905)

Bradycellus (Triliarthrus) georgei Lindroth, 1968

Bradycellus (Triliarthrus) kirbyi (Horn, 1883)

Bradycellus (Triliarthrus) lugubris (LeConte, 1847)

Amerinus linearis (LeConte, 1863)

Dicheirotrichus (Oreoxenus) mannerheimii mannerheimii (Sahlberg, 1844)

Dicheirotrichus (Trichocellus) cognatus (Gyllenhal, 1827)

Acupalpus (Acupalpus) canadensis Casey, 1924

Acupalpus (Acupalpus) carus (LeConte, 1863)

Acupalpus (Acupalpus) hydropicus (LeConte, 1863)

Acupalpus (Acupalpus) meridianus (Linnaeus, 1760)

Acupalpus (Acupalpus) nanellus Casey, 1914

Acupalpus (Acupalpus) pumilus Lindroth, 1968

Acupalpus (Tachistodes) indistinctus Dejean, 1831

Acupalpus (Tachistodes) partiarius (Say, 1823)

Acupalpus (Tachistodes) pauperculus Dejean, 1829

Acupalpus (Tachistodes) testaceus Dejean, 1829

Acupalpus (Anthracus) punctulatus Hatch, 1953

Acupalpus (Anthracus) tener (LeConte, 1857)

Philodes (Philodes) alternans (LeConte, 1853)

Philodes (Goniolophus) flavilimbus (LeConte, 1869)

Philodes (Goniolophus) longulus (Dejean, 1829)

Philodes (Goniolophus) rectangulus (Chaudoir, 1868)

Pogonodaptus mexicanus (Bates, 1878)

Polpochila (Phymatocephalus) capitata (Chaudoir, 1852)

Polpochila (Phymatocephalus) erro (LeConte, 1854)

Polpochila (Polpochila) rotundicollis Bates, 1882

Piosoma setosum LeConte, 1847

Euryderus grossus (Say, 1830)

Ophonus (Metophonus) puncticeps Stephens, 1828

Ophonus (Metophonus) rufibarbis (Fabricius, 1792)

Harpalus (Pseudoophonus) actiosus Casey, 1914

Harpalus (Pseudoophonus) compar LeConte, 1847

Harpalus (Pseudoophonus) erythropus Dejean, 1829

Harpalus (Pseudoophonus) faunus Say, 1823

Harpalus (Pseudoophonus) hatchi Ball & Anderson, 1962

Harpalus (Pseudoophonus) liobasis Chaudoir, 1868

Harpalus (Pseudoophonus) paratus Casey, 1924

Harpalus (Pseudoophonus) pensylvanicus (DeGeer, 1774)

Harpalus (Pseudoophonus) poncei Will, 2002

Harpalus (Pseudoophonus) protractus Casey, 1914

Harpalus (Pseudoophonus) rufipes (DeGeer, 1774)

Harpalus (Pseudoophonus) texanus Casey, 1914

Harpalus (Pseudoophonus) vagans LeConte, 1865

Harpalus (Megapangus) caliginosus (Fabricius, 1775)

Harpalus (Megapangus) katiae Battoni, 1985

Harpalus (Plectralidus) erraticus Say, 1823

Harpalus (Plectralidus) retractus LeConte, 1863

Harpalus (Opadius) animosus Casey, 1924

Harpalus (Opadius) apache Kataev, 2010

Harpalus (Opadius) cordatus (LeConte, 1853)

Harpalus (Opadius) cordifer Notman, 1919

Harpalus (Opadius) desertus LeConte, 1859

Harpalus (Opadius) fraternus LeConte, 1852

Harpalus (Opadius) fulvilabris Mannerheim, 1853

Harpalus (Opadius) gravis LeConte, 1858

Harpalus (Opadius) indianus Csiki, 1932

Harpalus (Opadius) indigens Casey, 1924

Harpalus (Opadius) laevipes Zetterstedt, 1828

Harpalus (Opadius) laticeps LeConte, 1850

Harpalus (Opadius) lewisii LeConte, 1865

Harpalus (Opadius) megacephalus LeConte, 1847

Harpalus (Opadius) nigritarsis Sahlberg, 1827

Harpalus (Opadius) providens Casey, 1914

Harpalus (Opadius) reversus Casey, 1924

Harpalus (Opadius) spadiceus Dejean, 1829

Harpalus (Opadius) ventralis LeConte, 1847

Harpalus (Harpalus) affinis (Schrank, 1781)

Harpalus (Harpalus) amputatus amputatus Say, 1830

Harpalus (Harpalus) atrichatus Hatch, 1949

Harpalus (Harpalus) balli Noonan, 1991

Harpalus (Harpalus) cautus Dejean, 1829

Harpalus (Harpalus) ellipsis LeConte, 1847

Harpalus (Harpalus) herbivagus Say, 1823

Harpalus (Harpalus) innocuus LeConte, 1863

Harpalus (Harpalus) martini Van Dyke, 1926

Harpalus (Harpalus) obnixus Casey, 1924

Harpalus (Harpalus) ochropus Kirby, 1837

Harpalus (Harpalus) opacipennis (Haldeman, 1843)

Harpalus (Harpalus) plenalis Casey, 1914

Harpalus (Harpalus) rubripes (Duftschmid, 1812)

Harpalus (Harpalus) solitaris Dejean, 1829

Harpalus (Harpalus) somnulentus Dejean, 1829

Harpalus (Harpalus) vittatus alaskensis Lindroth, 1968

Harpalus (Glanodes) cohni Ball, 1972

Harpalus (Glanodes) corpulentus (Casey, 1914)

Harpalus (Glanodes) huachuca Ball, 1972

Harpalus (Glanodes) obliquus Horn, 1880

Harpalus (Glanodes) puncticeps (Casey, 1914)

Harpalus (Glanodes) stephani Ball, 1972

Harpalus (Harpalobius) fuscipalpis Sturm, 1818

Harpalobrachys leiroides (Motschulsky, 1844)

Hartonymus alternatus (LeConte, 1863)

Hartonymus hoodi Casey, 1914

Amblygnathus evansi Ball & Maddison, 1987

Amblygnathus iripennis (Say, 1823)

Amblygnathus mexicanus Bates, 1882

Amblygnathus subtinctus (LeConte, 1867)

Athrostictus punctatulus (Putzeys, 1878)

Selenophorus (Celiamorphus) adjunctus (Casey, 1914)

Selenophorus (Celiamorphus) contractus (Casey, 1914)

Selenophorus (Celiamorphus) discopunctatus Dejean, 1829

Selenophorus (Celiamorphus) ellipticus Dejean, 1829

Selenophorus (Celiamorphus) fossulatus Dejean, 1829

Selenophorus (Celiamorphus) granarius Dejean, 1829

Selenophorus (Celiamorphus) municeps (Casey, 1924)

Selenophorus (Celiamorphus) nanulus (Casey, 1924)

Selenophorus (Celiamorphus) subtropicus (Casey, 1924)

Selenophorus (Selenophorus) aeneopiceus Casey, 1884

Selenophorus (Selenophorus) blanchardi Manee, 1915

Selenophorus (Selenophorus) chaparralus Purrington, 2000

Selenophorus (Selenophorus) concinnus Schaeffer, 1910

Selenophorus (Selenophorus) cupreolus Casey, 1914

Selenophorus (Selenophorus) discoderoides Schaeffer, 1910

Selenophorus (Selenophorus) elongatus (LeConte, 1847)

Selenophorus (Selenophorus) famulus Casey, 1914

Selenophorus (Selenophorus) fatuus LeConte, 1863

Selenophorus (Selenophorus) gagatinus Dejean, 1829

Selenophorus (Selenophorus) houstoni Casey, 1914

Selenophorus (Selenophorus) hylacis (Say, 1823)

Selenophorus (Selenophorus) implicans Casey, 1914

Selenophorus (Selenophorus) integer (Fabricius, 1798)

Selenophorus (Selenophorus) laesus (LeConte, 1858)

Selenophorus (Selenophorus) maritimus Casey, 1914

Selenophorus (Selenophorus) opalinus (LeConte, 1863)

Selenophorus (Selenophorus) otiosus Casey, 1914

Selenophorus (Selenophorus) palliatus (Fabricius, 1798)

Selenophorus (Selenophorus) parumpunctatus Dejean, 1829

Selenophorus (Selenophorus) pedicularius Dejean, 1829

Selenophorus (Selenophorus) planipennis LeConte, 1847

Selenophorus (Selenophorus) riparius Casey, 1914

Selenophorus (Selenophorus) schaefferi Csiki, 1932

Selenophorus (Selenophorus) scolopaceus Casey, 1914

Selenophorus (Selenophorus) sinuaticollis Notman, 1922

Selenophorus (Selenophorus) striatopunctatus Putzeys, 1878

Selenophorus (Selenophorus) trepidus (Casey, 1924)

Selenophorus breviusculus Horn, 1880

Discoderus aequalis Casey, 1914

Discoderus amoenus LeConte, 1863

Discoderus congruens Casey, 1914

Discoderus cordicollis Horn, 1891

Discoderus crassicollis Horn, 1891

Discoderus dallasensis Casey, 1924

Discoderus impotens (LeConte, 1858)

Discoderus longicollis Casey, 1914

Discoderus obsidianus Casey, 1914

Discoderus papagonis Casey, 1924

Discoderus parallelus (Haldeman, 1843)

Discoderus parilis (Casey, 1914)

Discoderus peregrinus Casey, 1924

Discoderus pinguis Casey, 1884

Discoderus robustus piceus Casey, 1914

Discoderus robustus robustus Horn, 1883

Discoderus subviolaceus Casey, 1914

Discoderus symbolicus Casey, 1914

Discoderus tenebrosus (LeConte, 1847)

Discoderus texanus Casey, 1924

Stenomorphus californicus californicus (Ménétriés, 1843)

Stenomorphus californicus rufipes LeConte, 1858

Stenomorphus convexior Notman, 1922

Stenomorphus sinaloae Darlington, 1936

Trichotichnus (Trichotichnus) dichrous (Dejean, 1829)

Trichotichnus (Trichotichnus) vulpeculus (Say, 1823)

Trichotichnus (Iridessus) autumnalis (Say, 1823)

Trichotichnus (Iridessus) fulgens (Csiki, 1932)

Aztecarpalus schaefferi Ball, 1970

Cratacanthus dubius (Palisot de Beauvois, 1811)


Pseudamara arenaria (LeConte, 1847)

Calathus (Calathus) fuscipes (Goeze, 1777)

Calathus (Neocalathus) calceus Ball & Nègre, 1972

Calathus (Neocalathus) gregarius (Say, 1823)

Calathus (Neocalathus) ingratus Dejean, 1828

Calathus (Neocalathus) opaculus LeConte, 1854

Calathus (Neocalathus) peropacus Casey, 1920

Calathus (Neocalathus) ruficollis grandicollis Casey, 1920

Calathus (Neocalathus) ruficollis ignicollis Casey, 1920

Calathus (Neocalathus) ruficollis ruficollis Dejean, 1828

Calathus (Acalathus) advena (LeConte, 1846)

Synuchus dubius (LeConte, 1854)

Synuchus impunctatus (Say, 1823)

Laemostenus (Laemostenus) complanatus (Dejean, 1828)

Laemostenus (Pristonychus) terricola terricola (Herbst, 1784)


Olisthopus brevicornis Casey, 1913

Olisthopus filicornis Casey, 1913

Olisthopus innuens Casey, 1913

Olisthopus iterans Casey, 1913

Olisthopus micans LeConte, 1846

Olisthopus parmatus (Say, 1823)

Olisthopus pusio Casey, 1913

Elliptoleus acutesculptus Bates, 1882

Sericoda bembidioides Kirby, 1837

Sericoda bogemannii (Gyllenhal, 1813)

Sericoda obsoleta (Say, 1823)

Sericoda quadripunctata (DeGeer, 1774)

Tetraleucus picticornis (Newman, 1844)

Anchomenus (Anchomenus) aeneolus (LeConte, 1854)

Anchomenus (Anchomenus) funebris (LeConte, 1854)

Anchomenus (Anchomenus) quadratus (LeConte, 1854)

Rhadine albamontana Dajoz, 1998

Rhadine anthicoides Casey, 1913

Rhadine austinica Barr, 1974

Rhadine babcocki (Barr, 1960)

Rhadine balesi (Gray, 1937)

Rhadine bullis Reddell & Cokendolpher, 2004

Rhadine caudata (LeConte, 1863)

Rhadine constricta Casey, 1913

Rhadine dissecta (LeConte, 1863)

Rhadine exilis (Barr & Lawrence, 1960)

Rhadine grubbsi Reddell & Dupérré, 2009

Rhadine howdeni (Barr & Lawrence, 1960)

Rhadine infernalis ewersi (Barr, 1960)

Rhadine infernalis infernalis (Barr & Lawrence, 1960)

Rhadine insolita Barr, 1974

Rhadine ivyi Reddell & Cokendolpher, 2004

Rhadine jejuna (LeConte, 1878)

Rhadine koepkei koepkei (Barr, 1960)

Rhadine koepkei privata Barr, 1974

Rhadine lanei (Gray, 1937)

Rhadine larvalis LeConte, 1846

Rhadine lindrothi Barr, 1965

Rhadine longiceps Van Dyke, 1949

Rhadine longicollis Benedict, 1927

Rhadine longipes Casey, 1913

Rhadine myrmecodes (Horn, 1892)

Rhadine nivalis (Horn, 1881)

Rhadine noctivaga Barr, 1974

Rhadine ozarkensis Sanderson & Miller, 1941

Rhadine perlevis Casey, 1913

Rhadine persephone Barr, 1974

Rhadine pertenuis Casey, 1920

Rhadine reyesi Reddell & Cokendolpher, 2001

Rhadine rossi Van Dyke, 1949

Rhadine rubra (Barr, 1960)

Rhadine russelli Barr, 1974

Rhadine specum crinicollis Barr, 1974

Rhadine specum gentilis Barr, 1974

Rhadine specum specum (Barr, 1960)

Rhadine sprousei Reddell & Cokendolpher, 2004

Rhadine sublustris Casey, 1913

Rhadine subterranea mitchelli Barr, 1974

Rhadine subterranea subterranea (Van Dyke, 1919)

Rhadine tenebrosa mckenziei Barr, 1974

Rhadine tenebrosa tenebrosa (Barr, 1960)

Rhadine testacea Casey, 1920

Rhadine umbra Casey, 1913

Mexisphodrus valverdensis Barr, 1982

Tanystoma cuyama Liebherr, 1985

Tanystoma maculicolle (Dejean, 1828)

Tanystoma striatum (Dejean, 1828)

Tanystoma sulcatum (Dejean, 1828)

Paranchus albipes (Fabricius, 1794)

Oxypselaphus pusillus (LeConte, 1854)

Agonum (Platynomicrus) ferruginosum (Dejean, 1828)

Agonum (Platynomicrus) nigriceps LeConte, 1846

Agonum (Europhilus) anchomenoides Randall, 1838

Agonum (Europhilus) canadense Goulet, 1969

Agonum (Europhilus) consimile (Gyllenhal, 1810)

Agonum (Europhilus) darlingtoni Lindroth, 1954

Agonum (Europhilus) exaratum (Mannerheim, 1853)

Agonum (Europhilus) galvestonicum (Casey, 1920)

Agonum (Europhilus) gratiosum (Mannerheim, 1853)

Agonum (Europhilus) limbatum Motschulsky, 1845

Agonum (Europhilus) lutulentum (LeConte, 1854)

Agonum (Europhilus) palustre Goulet, 1969

Agonum (Europhilus) picicornoides Lindroth, 1966

Agonum (Europhilus) retractum LeConte, 1846

Agonum (Europhilus) simile Kirby, 1837

Agonum (Europhilus) sordens Kirby, 1837

Agonum (Europhilus) superioris Lindroth, 1966

Agonum (Europhilus) thoreyi Dejean, 1828

Agonum (Agonum) bicolor (Dejean, 1828)

Agonum (Agonum) muelleri (Herbst, 1784)

Agonum (Agonum) piceolum (LeConte, 1879)

Agonum (Agonum) placidum (Say, 1823)

Agonum (Olisares) aeruginosum Dejean, 1828

Agonum (Olisares) affine Kirby, 1837

Agonum (Olisares) albicrus Dejean, 1828

Agonum (Olisares) anthracinum Dejean, 1831

Agonum (Olisares) basale LeConte, 1846

Agonum (Olisares) belleri (Hatch, 1933)

Agonum (Olisares) brevicolle Dejean, 1828

Agonum (Olisares) collare (Say, 1830)

Agonum (Olisares) corvus (LeConte, 1860)

Agonum (Olisares) crenistriatum (LeConte, 1863)

Agonum (Olisares) crenulatum (LeConte, 1854)

Agonum (Olisares) cupreum Dejean, 1831

Agonum (Olisares) cupripenne (Say, 1823)

Agonum (Olisares) cyanopis (Bates, 1882)

Agonum (Olisares) cyclifer (Bates, 1884)

Agonum (Olisares) deceptivum (LeConte, 1879)

Agonum (Olisares) decorum (Say, 1823)

Agonum (Olisares) deplanatum Ménétriés, 1843

Agonum (Olisares) elongatulum (Dejean, 1828)

Agonum (Olisares) errans (Say, 1823)

Agonum (Olisares) excavatum Dejean, 1828

Agonum (Olisares) extensicolle (Say, 1823)

Agonum (Olisares) extimum Liebherr, 1986

Agonum (Olisares) ferreum Haldeman, 1843

Agonum (Olisares) fidele Casey, 1920

Agonum (Olisares) fossiger Dejean, 1828

Agonum (Olisares) harrisii LeConte, 1846

Agonum (Olisares) imitans (Notman, 1919)

Agonum (Olisares) melanarium Dejean, 1828

Agonum (Olisares) metallescens (LeConte, 1854)

Agonum (Olisares) moerens Dejean, 1828

Agonum (Olisares) muiri Liebherr, 1984

Agonum (Olisares) mutatum (Gemminger & Harold, 1868)

Agonum (Olisares) nutans (Say, 1823)

Agonum (Olisares) octopunctatum (Fabricius, 1798)

Agonum (Olisares) pacificum Casey, 1920

Agonum (Olisares) pallipes (Fabricius, 1787)

Agonum (Olisares) parextimum Liebherr, 1986

Agonum (Olisares) propinquum (Gemminger & Harold, 1868)

Agonum (Olisares) punctiforme (Say, 1823)

Agonum (Olisares) quadrimaculatum (Horn, 1885)

Agonum (Olisares) quinquepunctatum Motschulsky, 1844

Agonum (Olisares) rigidulum (Casey, 1920)

Agonum (Olisares) rufipes Dejean, 1828

Agonum (Olisares) striatopunctatum Dejean, 1828

Agonum (Olisares) sulcipenne (Horn, 1881)

Agonum (Olisares) suturale Say, 1830

Agonum (Olisares) tenue (LeConte, 1854)

Agonum (Olisares) texanum (LeConte, 1878)

Agonum (Olisares) trigeminum Lindroth, 1954

Platynus (Microplatynus) agilis LeConte, 1863

Platynus (Microplatynus) pecki Barr, 1982

Platynus (Platynus) brunneomarginatus (Mannerheim, 1843)

Platynus (Platynus) daviesi Bousquet, 2012

Platynus (Platynus) decentis (Say, 1823)

Platynus (Platynus) indecentis Liebherr & Will, 1996

Platynus (Platynus) opaculus LeConte, 1863

Platynus (Platynus) ovipennis (Mannerheim, 1843)

Platynus (Platynus) parmarginatus Hamilton, 1893

Platynus (Platynus) tenuicollis (LeConte, 1846)

Platynus (Platynus) trifoveolatus Beutenmüller, 1903

Platynus (Batenus) angustatus Dejean, 1828

Platynus (Batenus) cincticollis (Say, 1823)

Platynus (Batenus) hypolithos (Say, 1823)

Platynus (Batenus) mannerheimii (Dejean, 1828)

Platynus (Batenus) prognathus Van Dyke, 1926

Platynus (Glyptolenopsis) ovatulus (Bates, 1884)

Platynus (Trapezodera) cohni Liebherr & Will, 1996

Platynus (Dyscolus) cazieri Liebherr & Will, 1996

Platynus (Dyscolus) falli (Darlington, 1936)

Platynus (Dyscolus) lyratus (Chaudoir, 1879)

Platynus (Dyscolus) megalops (Bates, 1882)

Platynus (Dyscolus) rufiventris (Van Dyke, 1926)

Metacolpodes buchanani (Hope, 1831)


Perigona (Trechicus) nigriceps (Dejean, 1831)

Perigona (Trechicus) pallipennis (LeConte, 1853)


Atranus pubescens (Dejean, 1828)


Anchonoderus quadrinotatus Horn, 1878

Anchonoderus schaefferi Liebke, 1928

Lachnophorus elegantulus Mannerheim, 1843

Euphorticus occidentalis Horn, 1891

Euphorticus pubescens (Dejean, 1831)

Calybe (Ega) laetula (LeConte, 1851)

Calybe (Ega) sallei (Chevrolat, 1839)

Eucaerus (Eucaerus) varicornis LeConte, 1853


Pentagonica bicolor (LeConte, 1863)

Pentagonica felix Bell, 1987

Pentagonica flavipes flavipes (LeConte, 1853)

Pentagonica marshalli Mateu, 1995

Pentagonica nigricornis Darlington, 1934

Pentagonica picticornis Bates, 1883


Colliuris (Mimocasnonia) pilatei (Chaudoir, 1848)

Colliuris (Cosnania) lengi (Schaeffer, 1910)

Colliuris (Cosnania) pensylvanica (Linnaeus, 1758)

Colliuris (Calocolliuris) caymanensis Darlington, 1947

Colliuris (Calocolliuris) lioptera (Bates, 1891)

Colliuris (Calocolliuris) ludoviciana (Sallé, 1849)


Leptotrachelus depressus Blatchley, 1923

Leptotrachelus dorsalis (Fabricius, 1801)

Leptotrachelus pallidulus Motschulsky, 1864


Tetragonoderus (Crossonychus) fasciatus (Haldeman, 1843)

Tetragonoderus (Crossonychus) intersectus (Germar, 1824)

Tetragonoderus (Crossonychus) laevigatus Chaudoir, 1876

Tetragonoderus (Crossonychus) latipennis LeConte, 1874

Tetragonoderus (Crossonychus) pallidus Horn, 1869


Mochtherus tetraspilotus (Macleay, 1825)

Phloeoxena (Oenaphelox) signata (Dejean, 1825)

Eucheila (Inna) boyeri (Solier, 1835)

Somotrichus unifasciatus (Dejean, 1831)

Coptodera (Coptodera) aerata Dejean, 1825

Coptodera (Coptodera) brunnea Shpeley & Ball, 1994

Coptodera (Coptodera) festiva Dejean, 1825

Coptodera (Coptodera) nitidula (Buquet, 1835)

Coptodera (Coptodera) picea Dejean, 1826

Cymindis (Tarulus) americana Dejean, 1826

Cymindis (Tarulus) arizonensis Schaeffer, 1910

Cymindis (Tarulus) borealis LeConte, 1863

Cymindis (Tarulus) californica Horn, 1895

Cymindis (Tarulus) cribricollis Dejean, 1831

Cymindis (Tarulus) elegans LeConte, 1846

Cymindis (Tarulus) evanescens Casey, 1913

Cymindis (Tarulus) interior Lindroth, 1969

Cymindis (Tarulus) laticollis Say, 1830

Cymindis (Tarulus) neglecta Haldeman, 1843

Cymindis (Tarulus) pilosa Say, 1823

Cymindis (Tarulus) planipennis LeConte, 1863

Cymindis (Tarulus) seriata Hatch, 1953

Cymindis (Tarulus) unicolor Kirby, 1837

Cymindis (Tarulus) uniseriata Bates, 1884

Cymindis (Tarulus) vaporariorum (Linnaeus, 1758)

Cymindis (Pinacodera) abbreviata (Casey, 1920)

Cymindis (Pinacodera) ampliata (Casey, 1920)

Cymindis (Pinacodera) atripennis (Casey, 1920)

Cymindis (Pinacodera) blanda Casey, 1913

Cymindis (Pinacodera) complanata Dejean, 1826

Cymindis (Pinacodera) limbata Dejean, 1831

Cymindis (Pinacodera) obscura (Casey, 1920)

Cymindis (Pinacodera) platicollis (Say, 1823)

Cymindis (Pinacodera) punctifera (LeConte, 1884)

Cymindis (Pinacodera) punctigera LeConte, 1851

Cymindis (Pinacodera) subcarinata (Casey, 1920)

Apenes (Apenes) angustata Schwarz, 1878

Apenes (Apenes) coriacea (Chevrolat, 1863)

Apenes (Apenes) hilariola Bates, 1891

Apenes (Apenes) lucidula lucidula (Dejean, 1831)

Apenes (Apenes) nebulosa LeConte, 1867

Apenes (Apenes) opaca LeConte, 1851

Apenes (Apenes) pallidipes (Chevrolat, 1836)

Apenes (Apenes) parallela parallela (Dejean, 1825)

Apenes (Apenes) sinuata (Say, 1823)

Dromius (Dromius) fenestratus (Fabricius, 1794)

Dromius (Dromius) piceus Dejean, 1831

Philorhizus atriceps (LeConte, 1880)

Philorhizus melanocephalus (Dejean, 1825)

Microlestes brevilobus brevilobus Lindroth, 1969

Microlestes curtipennis (Casey, 1920)

Microlestes lindrothi Mateu, 1995

Microlestes linearis (LeConte, 1851)

Microlestes lucidus lucidus (LeConte, 1851)

Microlestes major Lindroth, 1969

Microlestes nigrinus (Mannerheim, 1843)

Microlestes pusio (LeConte, 1863)

Apristus actuosus Casey, 1920

Apristus agitatus Casey, 1920

Apristus cephalus Casey, 1920

Apristus constrictus Casey, 1920

Apristus latens (LeConte, 1846)

Apristus laticollis LeConte, 1851

Apristus liratus Casey, 1920

Apristus nevadensis Casey, 1920

Apristus pugetanus Casey, 1920

Apristus subdeletus Casey, 1920

Apristus subsulcatus (Dejean, 1826)

Apristus thoracicus Casey, 1920

Apristus tuckeri Casey, 1920

Syntomus americanus (Dejean, 1831)

Axinopalpus biplagiatus (Dejean, 1825)

Axinopalpus denticulatus Hatch, 1949

Axinopalpus fusciceps LeConte, 1851

Axinopalpus illectus Casey, 1920

Axinopalpus pratti Hatch, 1949

Axinopalpus utahensis Tanner, 1928

Axinopalpus vittatus Hatch, 1949

Lebia (Loxopeza) atriceps LeConte, 1863

Lebia (Loxopeza) atriventris Say, 1823

Lebia (Loxopeza) deceptrix Madge, 1967

Lebia (Loxopeza) grandis Hentz, 1830

Lebia (Loxopeza) pimalis (Casey, 1920)

Lebia (Loxopeza) subdola Madge, 1967

Lebia (Loxopeza) subgrandis Madge, 1967

Lebia (Loxopeza) tricolor Say, 1823

Lebia (Polycheloma) lecontei Madge, 1967

Lebia (Lamprias) divisa LeConte, 1850

Lebia (Lebia) abdita Madge, 1967

Lebia (Lebia) abdominalis Chaudoir, 1843

Lebia (Lebia) analis Dejean, 1825

Lebia (Lebia) arizonica Schaeffer, 1910

Lebia (Lebia) bilineata Motschulsky, 1859

Lebia (Lebia) bitaeniata Chevrolat, 1834

Lebia (Lebia) bivittata (Fabricius, 1798)

Lebia (Lebia) bumeliae Schaeffer, 1910

Lebia (Lebia) calliope Bates, 1883

Lebia (Lebia) collaris Dejean, 1826

Lebia (Lebia) cyanipennis Dejean, 1831

Lebia (Lebia) esurialis Casey, 1920

Lebia (Lebia) fuscata Dejean, 1825

Lebia (Lebia) guttula LeConte, 1851

Lebia (Lebia) histrionica Bates, 1883

Lebia (Lebia) insulata Madge, 1967

Lebia (Lebia) lecta Horn, 1885

Lebia (Lebia) lobulata LeConte, 1863

Lebia (Lebia) marginicollis Dejean, 1825

Lebia (Lebia) miranda (Horn, 1872)

Lebia (Lebia) moesta LeConte, 1850

Lebia (Lebia) nigricapitata Madge, 1967

Lebia (Lebia) ornata Say, 1823

Lebia (Lebia) pectita Horn, 1885

Lebia (Lebia) perita Casey, 1920

Lebia (Lebia) perpallida Madge, 1967

Lebia (Lebia) pleuritica LeConte, 1846

Lebia (Lebia) pulchella Dejean, 1826

Lebia (Lebia) pumila Dejean, 1831

Lebia (Lebia) rufopleura Schaeffer, 1910

Lebia (Lebia) scalpta Bates, 1883

Lebia (Lebia) scapula Horn, 1885

Lebia (Lebia) solea Hentz, 1830

Lebia (Lebia) subrugosa Chaudoir, 1871

Lebia (Lebia) tuckeri (Casey, 1920)

Lebia (Lebia) viridipennis Dejean, 1826

Lebia (Lebia) viridis Say, 1823

Lebia (Lebia) vittata (Fabricius, 1777)

Hyboptera auxiliadora Erwin, 2004

Plochionus (Menidius) amandus Newman, 1840

Plochionus (Menidius) bicolor Notman, 1919

Plochionus (Menidius) discoideus LeConte, 1880

Plochionus (Menidius) timidus Haldeman, 1843

Plochionus (Plochionus) pallens (Fabricius, 1775)

Tecnophilus croceicollis croceicollis (Ménétriés, 1843)

Tecnophilus croceicollis peigani Larson, 1969

Tecnophilus pilatei Chaudoir, 1877

Calleida (Calleida) circumcincta Bates, 1883

Calleida (Calleida) decora (Fabricius, 1801)

Calleida (Calleida) fimbriata Bates, 1883

Calleida (Calleida) fulgida Dejean, 1831

Calleida (Calleida) obrieni Mateu, 1995

Calleida (Calleida) planulata LeConte, 1858

Calleida (Calleida) platynoides Horn, 1882

Calleida (Calleida) punctata LeConte, 1846

Calleida (Calleida) punctulata Chaudoir, 1848

Calleida (Calleida) purpurea (Say, 1823)

Calleida (Calleida) viridipennis (Say, 1823)

Philophuga caerulea Casey, 1913

Philophuga viridicollis (LeConte, 1846)

Philophuga viridis amoena (LeConte, 1846)

Philophuga viridis horni Chaudoir, 1877

Philophuga viridis klamathea Larson, 1969

Philophuga viridis viridis (Dejean, 1831)

Infernophilus castaneus (Horn, 1882)

Onota angulicollis (Reiche, 1842)

Onota floridana Horn, 1881

Cylindronotum aeneum Putzeys, 1845

Agra oblongopunctata oblongopunctata Chevrolat, 1836

Euproctinus (Neoeuproctus) abjectus (Bates, 1883)

Euproctinus (Neoeuproctus) balli Shpeley, 1986

Euproctinus (Neoeuproctus) trivittatus (LeConte, 1878)

Nemotarsus elegans LeConte, 1853

Nemotarsus rhombifer Bates, 1883


Zuphium americanum Dejean, 1831

Zuphium delectum Liebke, 1933

Zuphium longicolle LeConte, 1879

Zuphium magnum Schaeffer, 1910

Zuphium mexicanum Chaudoir, 1863

Zuphium pseudamericanum Mateu, 1981

Pseudaptinus (Pseudaptinus) lecontei (Dejean, 1831)

Pseudaptinus (Pseudaptinus) oviceps Van Dyke, 1926

Pseudaptinus (Pseudaptinus) tenuicollis (LeConte, 1851)

Pseudaptinus (Thalpius) cubanus (Chaudoir, 1877)

Pseudaptinus (Thalpius) deceptor Darlington, 1934

Pseudaptinus (Thalpius) dorsalis (Brullé, 1834)

Pseudaptinus (Thalpius) hoegei (Bates, 1883)

Pseudaptinus (Thalpius) horni (Chaudoir, 1872)

Pseudaptinus (Thalpius) microcephalus (Van Dyke, 1926)

Pseudaptinus (Thalpius) nobilis Liebke, 1934

Pseudaptinus (Thalpius) pygmaeus (Dejean, 1826)

Pseudaptinus (Thalpius) rufulus (LeConte, 1851)


Galerita (Progaleritina) atripes LeConte, 1858

Galerita (Progaleritina) bicolor (Drury, 1773)

Galerita (Progaleritina) forreri Bates, 1883

Galerita (Progaleritina) janus (Fabricius, 1792)

Galerita (Progaleritina) lecontei lecontei Dejean, 1831

Galerita (Progaleritina) mexicana Chaudoir, 1872

Galerita (Progaleritina) reichardti Ball & Nimmo, 1983

Galerita (Galerita) aequinoctialis Chaudoir, 1852


Helluomorphoides clairvillei (Dejean, 1831)

Helluomorphoides ferrugineus (LeConte, 1853)

Helluomorphoides latitarsis (Casey, 1913)

Helluomorphoides nigripennis (Dejean, 1831)

Helluomorphoides papago (Casey, 1913)

Helluomorphoides praeustus bicolor (Harris, 1828)

Helluomorphoides praeustus floridanus Ball, 1956

Helluomorphoides praeustus praeustus (Dejean, 1825)

Helluomorphoides texanus (LeConte, 1853)


Pseudomorpha (Pseudomorpha) alleni Van Dyke, 1953

Pseudomorpha (Pseudomorpha) alutacea Notman, 1925

Pseudomorpha (Pseudomorpha) augustata Horn, 1883

Pseudomorpha (Pseudomorpha) behrensi Horn, 1870

Pseudomorpha (Pseudomorpha) castanea Casey, 1909

Pseudomorpha (Pseudomorpha) champlaini Notman, 1925

Pseudomorpha (Pseudomorpha) consanguinea Notman, 1925

Pseudomorpha (Pseudomorpha) cronkhitei Horn, 1867

Pseudomorpha (Pseudomorpha) cylindrica Casey, 1889

Pseudomorpha (Pseudomorpha) excrucians Kirby, 1823

Pseudomorpha (Pseudomorpha) falli Notman, 1925

Pseudomorpha (Pseudomorpha) hubbardi Notman, 1925

Pseudomorpha (Pseudomorpha) parallela Van Dyke, 1943

Pseudomorpha (Pseudomorpha) schwarzi Notman, 1925

Pseudomorpha (Pseudomorpha) tenebroides Notman, 1925

Pseudomorpha (Pseudomorpha) vandykei Notman, 1925

Pseudomorpha (Pseudomorpha) vicina Notman, 1925

Pseudomorpha (Pseudomorpha) vindicata Notman, 1925

Catalogue of North American Geadephaga taxa
Family TRACHYPACHIDAE Thomson, 1857
Trachypachini C.G. Thomson, 1857: 5. Type genus: Trachypachus Motschulsky, 1844.

Six species in western North America (three species), South America (two species), and northern Eurasia (one species). The species are arrayed in two genera: Systolosoma Solier (two South American species) and Trachypachus (four species).

Genus Trachypachus Motschulsky, 1844
Trachypachus Motschulsky, 1844: 86. Type species: Blethisa zetterstedtii Gyllenhal, 1827 designated by Thomson (1859: 3). Etymology. Uncertain, possibly from the Greek trachelos (neck, by extension pronotum) contracted and pachys (thick), alluding to the convex pronotum (“corselet convexe, large”) or from the Greek trachys (uneven, rough) and pachys [masculine].
Trachypachys Gemminger and Harold, 1868a: 46. Unjustified emendation of Trachypachus Motschulsky, 1844.

Northern Hemisphere, with four species in the Nearctic (three species) and Palaearctic (one species) Regions.


Lindroth (1961a: 1-4) reviewed the North American species and discussed the structural differences between the three taxa.

Trachypachus gibbsii LeConte, 1861
Trachypachys gibbsii LeConte, 1861b: 339. Type locality: «east of Fort Colville [Washington]» (original citation). Syntype(s) in MCZ [# 85]. Etymology. The specific name honors the American geologist and ethnologist George Gibbs [1815-1873] who gathered zoological specimens for the Smithsonian while working for the Northwest Boundary Commission.
Trachypachus californicus Motschulsky, 1864: 194. Type locality: «Calif[ornie]» (original citation). Two syntypes, one listed as “corruptum, ” in ZMMU (Keleinikova 1976: 190). Synonymy established by Horn (1870a: 71).
Trachypachus alticola Casey, 1920: 144. Type locality: «Lake Tahoe [Placer County], California» (original citation). Lectotype (♂), designated by Lindroth (1975: 111), in USNM [# 46830]. Synonymy established by Van Dyke (1925: 112), confirmed by Lindroth (1961a: 3).

This species ranges from southern British Columbia (Lindroth 1961a: 4) to northwestern Montana (Russell 1968: 42; Edwards 1975: 48), south to the southern part of the Sierra Nevada (Lindroth 1961a: 4) and to the Coast Ranges (Fall 1901a: 39) in California.



Figure 4.

Trachypachus gibbsii LeConte. This species is a typical western element as are the other two North American trachypachids. Although superficially similar to some large Bembidion or small Amara and having the same ecological preferences, we now believe that these beetles, along with members of the related genus Systolosoma of South America, are not closely related to any groups of carabids.

Trachypachus inermis Motschulsky, 1850
Trachypachus inermis Motschulsky, 1850a: 16. Type locality: «California?» (original citation), herein restricted to Lake Tahoe, Placer County (see Casey 1920: 146, as Trachypachus specularis). Four syntypes in ZMMU (Keleinikova 1976: 201).
Trachypachus holmbergi Mannerheim, 1853: 119. Type locality: «ad ostia fl[umen] Kaktnu [= Kenai River] peninsulae Kenai [Alaska]» (original citation). Holotype [by monotypy] location unknown (possibly in ZMH). Synonymy established by LeConte (1857c: 31). Etymology. The specific name was proposed for the Finnish naturalist, geologist, and ethnographer Heinrich [Henrik] Johan Holmberg [1818-1864], who visited Russian America in 1850 and 1851 and collected insects.
Trachypachus oregonus Casey, 1920: 145. Type locality: «Oregon» (original citation). Lectotype (♀), designated by Lindroth (1975: 111), in USNM [# 46831]. Synonymy established by Van Dyke (1925: 112), confirmed by Lindroth (1961a: 1).
Trachypachus specularis Casey, 1920: 146. Type locality: «Lake Tahoe [Placer County], California» (original citation for the lectotype). Lectotype (♂), designated by Lindroth (1975: 111), in USNM [# 46832]. Synonymy established by Van Dyke (1925: 112), confirmed by Lindroth (1961a: 1).

The range of this species extends from the Kenai Peninsula in Alaska (Lindroth 1961a: 2) to northwestern Saskatchewan (Hooper 1980: 65), south to southern Colorado (Wickham 1902: 230; LeConte 1878a: 464; Bell 1971: 58), southern Utah (Garfield County, Foster F. Purrington pers. comm. 2011), and the Sierra Nevada and Coast Ranges in California (Fall 1901a: 39). One old specimen, simply labeled “Ks” is known (MCZ, collection LeConte).



Trachypachus slevini Van Dyke, 1925
Trachypachus slevini Van Dyke, 1925: 111. Type locality: «Olney [Clatsop County], near Astoria, Oregon» (original citation). Holotype (♀) in CAS [# 1616]. Etymology. The specific name honors Joseph Richard Slevin [1881-1957], curator of herpetology at the California Academy of Sciences from 1928 to 1957.

This species is known only from the western regions of Washington and Oregon (Lindroth 1961a: 4).



Family RHYSODIDAE Laporte, 1840
Rhysodites Laporte, 1840: 291. Type genus: Rhysodes Germar, 1822.

Worldwide, with about 355 species arrayed in six tribes: Clinidiini (about 135 species), Dhysorini (ten species), Leoglymmiini (one species), Medisorini (one species), Omoglymmiini (about 180 species), and Rhysodini (about 25 species). Over 90% of the species are found in the Southern Hemisphere.


Bell (1970) revised the North American, Middle American, and West Indies species and provided keys for their identification.

Tribe Clinidiini Bell and Bell, 1978
Clinidiina R.T. Bell and J.R. Bell, 1978: 59. Type genus: Clinidium Kirby, 1830.

Worldwide, with about 135 species arrayed in three genera: Clinidium (about 75 species), Grouvellina Bell and Bell (17 Madagascan species), and Rhyzodiastes Fairmaire (about 45 species). The vast majority of species are found in the Southern Hemisphere, with only 11 species (about 8% of the world fauna) occurring in the Northern Hemisphere.

Genus Clinidium Kirby, 1830
Clinidium Kirby, 1830: 6. Type species: Clinidium guildingii Kirby, 1830 by monotypy. Etymology (original). From the Greek clinidion (small couch), alluding to the body shape of the adult [neuter].

About 75 species in the Nearctic (six species), Neotropical (about 65 species), and Palaearctic (three species) Regions arrayed in four subgenera: Arctoclinidium (nine species), Clinidium s.str. (about 50 Neotropical species), Mexiclinidium Bell and Bell (11 Middle American species), and Tainoa Bell and Bell (four West Indian species).


Bell and Bell (1985) revised the species of the world and provided keys for their identification.

Subgenus Arctoclinidium Bell, 1970
Arctoclinidium R.T. Bell, 1970: 308. Type species: Rhysodes sculptilis Newman, 1838 by original designation. Etymology. From the Greek arctos (north) and the generic name Clinidium [q.v.] [neuter].

Northern Hemisphere, with nine species in North America (six species), Japan (one species), Caucasian region (one species), and southern Europe (one species).

Clinidium apertum allegheniense Bell and Bell, 1975
Clinidium allegheniense R.T. Bell and J.R. Bell, 1975: 65. Type locality: «Pittsburgh [Allegheny County], Pennsylvania» (original citation). Holotype (♂) in SMEK.

This subspecies is known only from southwestern Pennsylvania and the Black Mountains in western North Carolina (Bell and Bell 1985: 91). The record from “Ohio” (Bousquet and Larochelle 1993: 42) needs confirmation.



Clinidium apertum apertum Reitter, 1880
Clinidium apertum Reitter, 1880: 29. Type locality: «Himalaya» (original citation), which is incorrect (Bell and Bell 1985: 90); Cartersville, Bartow County, Georgia (see Bell and Bell 1975: 66, as Clinidium allegheniense georgicum), herein selected. Syntype(s) in NHMW (Bell and Bell 1985: 90).
Clinidium allegheniense georgicum R.T. Bell and J.R. Bell, 1975: 66. Type locality: «Cartersville [Bartow County], Georgia» (original citation). Holotype (♂) in USNM [# 73195]. Synonymy established by Bell and Bell (1978: 65).

This subspecies is known only from the type locality in northern Georgia (Bell and Bell 1985: 90).



Clinidium baldufi Bell, 1970
Clinidium baldufi R.T. Bell, 1970: 313. Type locality: «Dayton [La Salle County], Ill[inois]» (original citation). Holotype (♂) in MCZ [# 31748]. Etymology. The specific name was proposed for Walter Valentine Balduf [1889-1969], professor of entomology at the University of Illinois.

This species ranges from New Jersey to central Iowa, including southwestern Wisconsin (Messer 2010: 33), south to southern Mississippi (Bell and Bell 1985: 89) and northern Florida (Bell 1970: 313). Old specimens simply labeled from Nebraska, Kansas, Missouri, and Texas are known (Bell 1970: 313). The records from “Arkansas” and “Louisiana” (Bousquet and Larochelle 1993: 43) need confirmation.



Clinidium calcaratum LeConte, 1875
Clinidium calcaratum LeConte, 1875b: 164. Type locality: «Vancouver Island; Oregon» (original citation). Syntype(s) [3 originally cited] in MCZ [# 6831].

This species ranges from southern British Columbia, including Vancouver Island, south to Mendocino County in the Coast Ranges of California and Tuolumne County in the Sierra Nevada (Bell and Bell 1985: 84).



Clinidium rosenbergi Bell, 1970
Clinidium rosenbergi R.T. Bell, 1970: 315. Type locality: «Turkey Run State Park, Parke County, Indiana» (original citation). Holotype (♂) in MCZ [# 31749].

This species ranges from northwestern Pennsylvania to eastern Missouri, south to east-central Louisiana (West Feliciana Parish, Igor M. Sokolov pers. comm. 2009) and northern Alabama (Madison County, CMNH).



Clinidium sculptile (Newman, 1838)
Rhysodes sculptilis Newman, 1838b: 666. Type locality: «Wheeling [Ohio County], [West] Virginia» (lectotype label). Lectotype (♂), designated by Bell and Bell (1985: 92), in BMNH.
Rhysodes conjungens Germar, 1840a: 351 [nomen dubium]. Type locality: «Staaten Nordamerika’s» (original citation). Holotype [by monotypy] location unknown (possibly in ZMHB). Synonymy established by LeConte (1875b: 164).

The range of this species extends from the Catskills in southern New York to west-central Indiana (Bell and Bell 1985: 92), south to east-central Louisiana (West Feliciana Parish, Igor M. Sokolov pers. comm. 2009), northern Alabama (Bell and Bell 1985: 92), and the Florida Panhandle (Peck and Thomas 1998: 15). Specimens simply labeled from Texas are known (Bell and Bell 1985: 92). The record from central Illinois (Wolcott 1896: 235) needs confirmation.



Clinidium valentinei Bell, 1970
Clinidium valentinei R.T. Bell, 1970: 313. Type locality: «Gorgas, Walker County, Ala[bama]» (original citation). Holotype (♂) in OSUO.

This species seems to be confined to the Appalachian Mountains from southwestern Pennsylvania to north-central Alabama and northeastern Georgia (Bell and Bell 1985: 85).



Tribe Omoglymmiini Bell and Bell, 1978
Omoglymmiina R.T. Bell and J.R. Bell, 1978: 66. Type genus: Omoglymmius Ganglbauer, 1891.

Worldwide, with about 180 species arrayed in eight genera. The tribe is much more diverse, both in term of species and lineages, in Asia than anywhere else.

Genus Omoglymmius Ganglbauer, 1891
Omoglymmius Ganglbauer, 1891a: 533. Type species: Rhysodes germari Ganglbauer, 1891 by monotypy. Etymology. From the Greek omos (rough) and glymma (an engraved figure) [masculine].

About 150 species (Lorenz 2005: 158-159) in the Nearctic (two species), Australian, Oriental, Palaearctic (ten species, only one of them present in Europe), and Afrotropical (one species) Regions arrayed in 11 subgenera. More than 90% of the species are found in Asia.


Bell and Bell (1983) revised the species of the world and provided keys for their identification.

Subgenus Boreoglymmius Bell and Bell, 1983
Boreoglymmius R.T. Bell and J.R. Bell, 1983: 140. Type species: Rhysodes americanus Laporte, 1836 by original designation. Etymology. From the Greek bore (north) and the last two syllables of the generic name Omoglymmius [masculine].

Three species in North American (two species) and Japan (one species).

Omoglymmius americanus (Laporte, 1836)
Rhysodes exaratus Lepeletier and Audinet-Serville [in Latreille et al.], 1825: 308 [primary homonym of Rhysodes exaratus Dalman, 1823]. Type locality: «Amérique septentrionale» (original citation), herein restricted to Florence, Florence County, South Carolina (see Bell and Bell 1983: 145). Syntype(s) location unknown (possibly in MHNP).
Rhysodes americanus Laporte, 1836: 58. Replacement name for Rhysodes exaratus Lepeletier and Audinet-Serville, 1825.
Rhysodes aratus Newman, 1838b: 664. Type locality. «Alabama» (original citation). Syntype(s) location unknown (possibly in BMNH). Synonymy established by LeConte (1875b: 162).

This species ranges from central New York to eastern Minnesota, south to east-central Texas (Bell and Bell 1983: 145) and northern Florida (Peck and Thomas 1998: 15).



Figure 5.

Omoglymmius americanus (Laporte). This species is one of the seven rhysodid species-group taxa found in eastern North America. These species live in decaying wood, such as logs, stumps or roots, where they feed on slime molds and fungi. The carabids, on the other hand, are carnivorous, herbivorous, or omnivorous feeding on both animal and plant matters.

Omoglymmius hamatus (LeConte, 1875)
Rhysodes hamatus LeConte, 1875b: 163. Type locality: «California» (original citation), herein restricted to Big Trees, Calaveras County (see Bell and Bell 1983: 144). Syntype(s) in MCZ [# 6830].

This species ranges from northern Idaho and southeastern Washington south to southern California, including the Sierra Nevada, and southeastern Arizona (Bell and Bell 1983: 143-144). Old specimens without specific localities from British Columbia and Texas (Bell and Bell 1983: 144) are known.



Family CARABIDAE Latreille, 1802
Carabici Latreille, 1802: 80. Type genus: Carabus Linnaeus, 1758.
Subfamily NEBRIINAE Laporte, 1834
Nebriidae Laporte, 1834: 90. Type genus: Nebria Latreille, 1802.

About 665 species in the Nearctic (about 75 species), Neotropical (one South American species), and Palaearctic and northern parts of the Oriental (about 595 species) Regions. The species are arrayed in five tribes: Nebriini (about 600 species), Notiokasiini (one South American species), Notiophilini (about 55 species), Opisthiini (five species), and Pelophilini (two species).

Tribe Pelophilini Kavanaugh, 1996
Pelophilini Kavanaugh, 1996: 35. Type genus: Pelophila Dejean, 1821.

This tribe contains a single genus.

Genus Pelophila Dejean, 1821
Pelophila Dejean, 1821: 7. Type species: Carabus borealis Paykull, 1790 by monotypy. Etymology (see Dejean 1826: 263). From the Greek pelos (mud) and philos (beloved), alluding to the habitat of Pelophila borealis [feminine].

Northern Hemisphere, with one Holarctic species and one species endemic to northern North America.


Lindroth (1961a: 57-60) covered both species and discussed the structural differences between them.

Pelophila borealis (Paykull, 1790)
Carabus borealis Paykull, 1790: 51. Type locality: «Karungi Botniae occidentalis [Norrbotten County, Sweden]» (original citation). Syntype(s) probably in NRSS.
Pelophila gebleri Mannerheim, 1823: 38. Type locality: «Sibiria ad Barnaul [Altai Kray, Russia]» (original citation). One syntype in ZMH (Silfverberg 1987: 17). Synonymy established by Dejean (1833: 22).
Pelophila marginata Mannerheim, 1823: 39. Type locality: «peninsula Kamschatka ad portum St. Petri et Pauli [apparently the town of Petropawlowsk, see Lindroth (1961a: 74)]» (original citation). One syntype in ZMH (Silfverberg 1987: 20). Synonymy established by Dejean (1833: 22).
Pelophila eschscholtzii Mannerheim, 1823: 40. Type locality: «insula Unalaschka [Aleutian Islands, Alaska]» (original citation). Lectotype (♂), designated by Lindroth (1961a: 57), in ZMH. Synonymy established by Dejean (1833: 22), confirmed by Lindroth (1961a: 57).
Pelophila elongata Mannerheim, 1823: 41. Type locality: «peninsula Kamschatka ad portum St. Petri et Pauli [apparently the town of Petropawlowsk, see Lindroth (1961a: 74)]» (original citation). One syntype in ZMH (Silfverberg 1987: 15). Synonymy established by Dejean (1833: 22).
Pelophila borealis var. arctica Dejean, 1826: 265. Type locality not stated. Syntype(s) location unknown (possibly in MHNP). Synonymy established by Dejean (1833: 22).
Pelophila borealis var. dejeanii Dejean, 1826: 265. Type locality: «environs de Barnaoul, Sibérie [Altai Kray, Russia]» (original citation). Syntype(s) probably in MHNP. Synonymy established by Dejean (1833: 22).
Pelophila laevigata Motschulsky, 1844: 92. Type locality: «près de la redoute Yamyschevo non loin du fleuve Irtych [Siberia, Russia]» (original citation). Holotype [by monotypy] in ZMMU (Keleinikova 1976: 202). Synonymy established by Bänninger (1930: 101).
Pelophila californica Motschulsky, 1844: 93. Type locality: «Californie» (original citation), which is incorrect (Lindroth 1961a: 57). Syntype(s) location unknown (possibly in ZMMU though not listed in Keleinikova 1976). Synonymy established by Bänninger (1930: 101).
Pelophila ochotica R.F. Sahlberg, 1844: 17. Type locality: «in monte Morikan [Okhotsk, Khabarovsk Kray, Siberia, Russia]» (original citation). Holotype [by monotypy] location unknown (possibly in ZMUT). Synonymy established by Bänninger (1930: 101).
Pelophila angusticollis Motschulsky, 1860: 98. Type locality: «Kamtschatka» (original citation). Five syntypes in ZMMU (Keleinikova 1976: 186). Synonymy established by Shilenkov (1994: 9).
Pelophila ulkei G.H. Horn, 1870b: 105. Type locality: «Hudson’s Bay Territory» (original citation), restricted to «Gillam, Manit[oba]» by Lindroth (1961a: 58). Holotype [by monotypy] (♂) in MCZ (collection LeConte). Synonymy established by Lindroth (1961a: 57).
Pelophila shermani Casey, 1913: 45. Type locality: «West S[ain]t Modest[e], Labrador» (original citation). Five syntypes [5 originally cited] in USNM [# 46844]. Synonymy established, under the name Pelophila borealis ulkei Horn, by Bänninger (1930: 102), confirmed by Lindroth (1961a: 57). Etymology. The specific name was proposed for John Dempster Sherman [1872-1960], dealer of entomological and related books and periodicals.

This species is found from northern Europe to the Bering Sea coast (Farkač 2003: 98) and from Alaska, including the Aleutian and Kodiak Islands, to Newfoundland, south to the Abitibi region in western Quebec (Larochelle 1975: 98) [see Lindroth 1963a: Fig. 59]. Fossil remnants of this species, dated between about 14, 000 and 18, 100 years B.P., have been unearthed in southeastern Iowa (Baker et al. 1986: 96; Schwert 1992: 76) and southern Ontario (Morgan and Morgan 1981: 1107).


CAN: AB, BC, LB, MB, NF, NT, NU, ON, QC, SK, YT USA: AK – Holarctic

Figure 6.

Pelophila borealis (Paykull). This pelophiline is one of the 97 species-group taxa which are Holarctic and found naturally in both the Palaearctic and Nearctic Regions. Most of these species are confined to the arctic, subarctic or boreal regions but a few are found mainly in the temperate regions such as Dyschirius politus. Most biogeographers agree that these species have spread between the two continents through Beringia during the Quaternary period.

Pelophila rudis (LeConte, 1863)
Nebria rudis LeConte, 1863c: 3. Type locality: «Methy [Portage] [= Portage La Loche, northern Saskatchewan]» (original citation). Holotype [by monotypy] (♂) in MCZ [# 653]. Note. In the original description, LeConte indicated that he received “one specimen” from “Mr. Kennicott” collected at “Methy.” Lindroth (1961a: 59) placed the type locality in eastern Alberta. However, I was unable to find any such locality in Alberta, although there is a locality named “Metis” in northwestern Alberta. In the same paper, LeConte (1863c: 1) described his Cicindela hyperborea from specimen(s) received from “Mr. R. Kennicott” collected at “Methy Portage, Hudson’s Bay Territory.” I believe LeConte meant “Methy Portage” as the collection site for Pelophila rudis. Located in northern Saskatchewan, Methy(e) Portage is currently called Portage La Loche.

This species is known from scattered localities from western Newfoundland (Lindroth 1955a: 39) to the Fairbanks area in Alaska (David H. Kavanaugh pers. comm. 2009), south to central British Columbia (Lindroth 1961a: 59) [see Lindroth 1963a: Fig. 65]. Fossil remnants, dated between 14, 000 and 15, 500 years B.P., have been unearthed in central Iowa (Schwert 1992: 76).



Tribe Opisthiini Dupuis, 1912
Opisthiinae Dupuis, 1912: 1. Type genus: Opisthius Kirby, 1837.

Northern Hemisphere, with five species in North America (one species) and the Himalayas and China, including Taiwan (four species). The species are arrayed in two genera: Opisthius (one species) and Paropisthius Casey (four species).


Bousquet and Smetana (1996) reviewed the species and provided a key for their identification.

Genus Opisthius Kirby, 1837
Opisthius Kirby, 1837: 60. Type species: Opisthius richardsoni Kirby, 1837 by monotypy. Etymology. Uncertain, possibly from the Greek opisthen (behind) or opisthios (hinder) [masculine]. Bousquet and Smetana (1996: 218) suggested that the name possibly reflect Kirby’s assignment of Opisthius to follow Elaphrus in his paper.

One North American species.


The species was treated in Lindroth’s (1961a: 88-90) monograph on the Carabidae of Canada and Alaska.

Opisthius richardsoni Kirby, 1837
Opisthius richardsoni Kirby, 1837: 61. Type locality: «[probably] on an island of Lake Winnipeg» (original citation), which is incorrect (Lindroth 1961a: 89); «Medicine Hat, Al[ber]ta» selected by Lindroth (1961a: 89). Lectotype (♂), designated by Bousquet and Smetana (1996: 220), in BMNH. Etymology. The specific name was proposed for John Richardson [1787-1865], surgeon and naturalist to Sir John Franklin on two Arctic expeditions, 1819-1822 and 1825-1827.

The range of this species extends from central Saskatchewan to the Arctic Circle in central Alaska (Lindroth 1961a: 89-90), south to Tuolumne County in the Sierra Nevada of California (Dajoz 2007: 17) and north-central New Mexico (Taos County, CNC). The record from “Iowa” (Jaques and Redlinger 1946: 295) is probably based on a mislabeled specimen or a stray. Fossil remnants of this species from the late Wisconsinan age have been found in northeastern Illinois, north-central Iowa, south-central Minnesota, and northwestern Ontario (see Ashworth and Schwert 1991: 511); others from a Plio-Pleistocene sequence have been found in northwestern Greenland and Meighen Island (Böcher 1995: 18).



Tribe Nebriini Laporte, 1834
Nebriidae Laporte, 1834: 90. Type genus: Nebria Latreille, 1802.

About 600 species (Lorenz 2005: 116-124) in the Nearctic (about 60 species), Palaearctic, and northern parts of the Oriental Regions. The species are arrayed in four genera: Archastes Jedlička (27 Chinese species), Leistus (about 180 species), Nebria (including Oreonebria Daniel) (about 380 species), and Nippononebria (seven species).

Genus Leistus Frölich, 1799
Leistus Frölich, 1799: 9. Type species: Leistus testaceus Frölich, 1799 (= Carabus ferrugineus Linnaeus, 1758) designated by Daniel (1903: 171). Etymology. From the Greek leistos (to be carried off as booty, to be won by force) [masculine]. Note. Daniel (1903: 171) designated Carabus ferrugineus Linnaeus, 1758 as type species of Leistus Frölich, 1799, a species not originally included; however, since he listed the name in synonymy with Leistus testaceus Frölich, 1799, a species originally included, he is deemed to have designated the latter taxon as type species (ICZN 1999: Article 69.2.2).

Northern Hemisphere, with about 180 species (Lorenz 2005: 116-118) in the Nearctic (four species, one of them adventive) and Palaearctic (about 180 species) Regions. The species are arrayed in six subgenera: Evanoleistus Jedlička (about 95 Asian species), Leistus s.str. (about 40 species), Nebrileistus Bänninger (two species on Madeira and Canary Islands), Neoleistus (three species), Sardoleistus Perrault (one Mediterranean species), and Pogonophorus Latreille (about 35 Palaearctic species).

Subgenus Leistus Frölich, 1799
Leistus Frölich, 1799: 9. Type species: Leistus testaceus Frölich, 1799 (= Carabus ferrugineus Linnaeus, 1758) designated by Daniel (1903: 171).

About 40 Palaearctic species of which one is adventive in North America.


Larson (1978: 307-308) discussed the structural differences between the adventive species in North America and the three native species of the subgenus Neoleistus.

Leistus ferrugineus (Linnaeus, 1758)
Carabus ferrugineus Linnaeus, 1758: 415. Type locality: «Europa» (original citation). One possible syntype in LSL (Lindroth 1957b: 331).

This European species is adventive in North America where it is known only from near Saint John’s, Newfoundland (Larson 1978: 307). The first inventoried specimen collected on this continent was caught in 1977.


CAN: NF – Adventive

Subgenus Neoleistus Erwin, 1970
Neoleistus Erwin, 1970b: 112. Type species: Leistus ferruginosus Mannerheim, 1843 by original designation. Etymology. From the Greek prefix neo- (new) and the generic name Leistus [q.v.], probably alluding to the fact that these Leistus species inhabit the New World [masculine].

Three western North American species.


Erwin (1970b) revised the species and provided a key for their identification.

Taxonomic Note.

Perrault (1991a) added three species from the Far East (Leistus angulicollis Fairmaire, Leistus niger Gebler, and Leistus shenseensis Perrault) to this subgenus but Shilenkov (1999: 76) rejected this association and the Asian species are listed in the nominotypical subgenus by Farkač and Janata (2003: 81-82).

Leistus ferruginosus Mannerheim, 1843
Leistus ferrugineus Dejean, 1831: 569 [secondary homonym of Leistus ferrugineus (Linnaeus, 1758)]. Type locality: «détroit de Norfolk [= Sitka Sound, Baranof Island, Alaska], sur la côte nord-ouest de l’Amérique septentrionale» (original citation). Holotype [by monotypy] (♀) location unknown (possibly lost).
Leistus ferruginosus Mannerheim, 1843: 187. Type locality: «insula Sitkha [= Baranof Island, Alaska]» (original citation). Lectotype, designated by Lindroth (1961a: 56), in ZMH. Synonymy established by Mannerheim (1843: 188).
Leistus nigropiceus Casey, 1913: 45. Type locality: «Metlakatla, British Columbia» (original citation). Lectotype (♀), designated by Lindroth (1975: 111), in USNM [# 46843]. Synonymy established by Hatch (1949b: 115), confirmed by Lindroth (1954b: 121).

This species ranges from the Gulf of Alaska coast south to west-central Oregon, east to western Montana (Russell 1968: 44) [see Erwin 1970b: Fig. 7]. At least one specimen simply labeled from California is known (Erwin 1970b: 115).



Leistus longipennis Casey, 1920
Leistus longipennis Casey, 1920: 148. Type locality: «Humboldt Co[unty], California» (original citation). Holotype [by monotypy] (♂) in USNM [# 46842].

The range of this species is restricted to the extreme northwestern tip of Humboldt County in northern California (Erwin 1970b: 117) and southwestern Oregon (Curry County, James R. LaBonte pers. comm. 1992).



Leistus madmeridianus Erwin, 1970
Leistus madmeridianus Erwin, 1970b: 117. Type locality: «Jacoby Creek, 5.0 miles southeast of Arcata, Humboldt County, California» (original citation). Holotype (♂) in CAS [# 11312].

This species is known only from a few localities along the Pacific Coast in northern California [see Erwin 1970b: Fig. 7].



Genus Nippononebria Uéno, 1955
Nippononebria Uéno, 1955: 49. Type species: Nebria pusilla Uéno, 1955 by original designation. Etymology. From the English nippon (a Japanese name for Japan) and the generic name Nebria [q.v.], alluding to the country where these Nebria-like species known to Uéno lived [feminine].

Seven species in western North America (three species), Japan (three species), and Jilin Province in China (one species) arrayed in two subgenera: Nippononebria s.str. for the Japanese and Chinese species and Vancouveria for the Nearctic ones.

Taxonomic Note.

Kavanaugh (1995, 1996) regarded Nippononebria as the sister-group to Leistus while Ledoux and Roux (2005) listed Nippononebria and Vancouveria as subgenera of Nebria and suggested they form the sister-group to {Eonebria Semenov and Znojko + Sadonebria Ledoux and Roux}, a complex of 60 Palaearctic species. Because the North American students are used to Kavanaugh’s approach, the taxon is retained here as a distinct genus.

Subgenus Vancouveria Kavanaugh, 1995
Vancouveria Kavanaugh, 1995: 159. Type species: Nebria virescens Horn, 1870 by original designation. Etymology. From the geographic name Vancouver [feminine].

Three species in western North America.


Ledoux and Roux (2005: 712) reviewed the species and provided a key for their identification.

Nippononebria altisierrae (Kavanaugh, 1984)
Nebria altisierrae Kavanaugh, 1984: 160. Type locality: «Olmsted Point (2560 m), Yosemite National Park [Mariposa County], Sierra Nevada, California» (original citation). Holotype (♂) in CAS [# 14338].

This species occurs at high elevations in the Sierra Nevada of California, from El Dorado County south to Sequoia National Park [see Kavanaugh 1984: Fig. 31].



Nippononebria campbelli (Kavanaugh, 1984)
Nebria campbelli Kavanaugh, 1984: 161. Type locality: «Mount Baker (1460-1520 m), Cascade Range, Whatcom County, Washington» (original citation). Holotype (♂) in CAS [# 14339].

This species ranges from the Three Brothers Mountain in southern British Columbia south to Mount Baker in Washington [see Kavanaugh 1984: Fig. 31].



Nippononebria virescens (Horn, 1870)
Nebria virescens G.H. Horn, 1870b: 100. Type locality: «Vancouver [British Columbia]» (original citation). Holotype [by monotypy] in MCZ [# 652].
Nebria brevis Casey, 1913: 55. Type locality: «Corvallis [Benton County], Oregon» (original citation for the lectotype). Lectotype (♂), designated by Lindroth (1975: 112), in USNM [# 46862]. Synonymy established with doubt by Bänninger (1925: 261), confirmed by Lindroth (1961a: 76).

This species ranges from southwestern British Columbia, including Vancouver Island, south to the northern Sierra Nevada of California in Plumas County, east to the western edge of the Rockies in west-central Idaho (Kavanaugh 1978: 349).



Genus Nebria Latreille, 1802
Nebria Latreille, 1802: 89. Type species: Carabus brevicollis Fabricius, 1792 designated by Latreille (1810: 426). Etymology. According to Ledoux and Roux (2005: 29), the name came from the Greek nebrios (fawn), possibly alluding to the coloration of Nebria complanata, the first species cited by Latreille in the genus. However Latreille (1804: 275) stated that the name derived from nebrias which is part of the list of unknown fishes mentioned by the elders. According to Dalby (2003: 121), nebrias, cited in Aristotle and others, is perhaps the dogfish Scyliorhinus canicula [feminine].

About 380 species (535 species-group taxa) in the arctic, subarctic, boreal, and temperate areas of the Nearctic and Palaearctic (including northern Africa and the Canary Islands) Regions arrayed in 25 subgenera (Ledoux and Roux 2005: 76 excluding Nippononebria and Vancouveria). The North American fauna has 52 species (82 species-group taxa) placed in four subgenera.


Ledoux and Roux (2005) reviewed the species of the world and provided keys for the identification of the species. Lindroth’s (1961a) key included all North American species then known but many species-group taxa have been described subsequently by Kavanaugh (1979a, 1981b, 1984, 2008).

Taxonomic Note.

The species of Nebria (including Nippononebria) have been segregated in two main lineages by Ledoux and Roux (2005: 71-75), one (named Vetanebri) represented in the Palaearctic Region by 90 species and in the Nearctic Region by the three species of Vancouveria, the other one (Notanebri) containing about 290 species, 52 in the Nearctic and almost 240 in the Palaearctic.

Subgenus Boreonebria Jeannel, 1937
Boreonebria Jeannel, 1937b: 2. Type species: Carabus rufescens Strøm, 1768 (= Carabus gyllenhali Schönherr, 1806) by original designation. Etymology. From the Greek bore (north) and the generic name Nebria [q.v.], probably alluding to the northern ranges of the species of this taxon [feminine].

Thirty-one species (Ledoux and Roux 2005: 82) in North America (seven species) and Eurasia (26 species). Two species are Holarctic (Nebria frigida and Nebria nivalis).

[gyllenhali group]
Nebria crassicornis crassicornis Van Dyke, 1925
Nebria crassicornis Van Dyke, 1925: 121. Type locality: «Paradise Park, [Mount] Rainier National Park [Pierce County], Washington» (original citation). Holotype (♂) in CAS [# 1627].

This subspecies is confined to a small area of the Coast Ranges and Cascade Range in southwestern British Columbia and western Washington [see Kavanaugh 1988: Fig. 15].



Nebria crassicornis intermedia Van Dyke, 1949
Nebria intermedia Van Dyke, 1949a: 49. Type locality: «Logan Pass, Glacier National Park [Flathead County], Montana» (original citation). Holotype (♂) in CAS [# 6008].

This subspecies ranges from northern British Columbia south to northeastern Oregon and southern Utah, east to northwestern Wyoming and central Utah [see Kavanaugh 1988: Fig. 15].



Nebria frigida Sahlberg, 1844
Nebria frigida R.F. Sahlberg, 1844: 11. Type locality: «monte Morikan cepi [Okhotsk, Khabarovsk Kray, Siberia, Russia]» (original citation). Lectotype (♀), designated by Lindroth (1961a: 81), in ZMUT.
Nebria viridis G.H. Horn, 1870b: 101. Type locality: «St. Michaels [= Saint Michael on south coast of Norton Sound], Alaska» (original citation). Lectotype (♂), designated by Kavanaugh (1979a: 116), in MCZ [# 34044]. Synonymy established by Lindroth (1961a: 81).
Nebria parvula J.R. Sahlberg, 1885b: 47. Type locality: Port Clarence, Alaska (inferred from title of the paper). Holotype [by monotypy; designated lectotype by Lindroth (1961a: 81)] (♂) in NRSS. Synonymy established, under the name Nebria viridis Horn, by Van Dyke (1924a: 5), confirmed by Lindroth (1961a: 81).
Nebria reducta Casey, 1920: 150. Type locality: «S[ain]t Paul Island, Alaska» (original citation), which is incorrect according to Lindroth (1961a: 24, 81). Lectotype (♀), designated by Lindroth (1975: 112), in USNM [# 46866]. Synonymy established, under the name Nebria viridis Horn, by Van Dyke (1924a: 5), confirmed by Lindroth (1961a: 81).

This Holarctic species is found in eastern Siberia (Farkač and Janata 2003: 88) and from the Alaskan Coast Range to the Anderson River Delta in northern Northwest Territories, south to northern British Columbia (Kavanaugh 1978: 714-715).


CAN: BC, NT, YT USA: AK – Holarctic


This species is placed in its own group by Ledoux and Roux (2005: 82).

Nebria gyllenhali castanipes (Kirby, 1837)
Helobia castanipes Kirby, 1837: 20. Type locality: «Lat. 65° [= apparently region of Great Bear Lake, Northwest Territories]» (original citation), restricted to «Nipigon, Ont[ario]» by Lindroth (1961a: 78). Lectotype (♀), designated by Kavanaugh (1979a: 111), in BMNH.
Nebria moesta LeConte, 1850: 209. Type locality: Lake Superior (inferred from title of the paper). Lectotype (♂), designated by Kavanaugh (1979a: 114), in MCZ [# 645]. Synonymy established by LeConte (1873b: 322), confirmed by Lindroth (1954b: 121).
Nebria elias Motschulsky, 1866: 276 [nomen dubium]. Type locality: «Amer[ique] rus[se]» (original citation). Syntype(s) lost (Keleinikova 1976: 196; Kavanaugh 1979a: 112). Synonymy established with doubt by Lindroth (1961a: 78).
Nebria labradorica Casey, 1920: 151. Type locality: «West S[ain]t Modest[e], Labrador» (original citation). Lectotype (♀), designated by Lindroth (1975: 112), in USNM [# 46855]. Synonymy established by Lindroth (1954b: 122).
Nebria prominens Casey, 1920: 151. Type locality: «M[oun]t Washington [Coos County], New Hampshire» (original citation). Lectotype (♀), designated by Lindroth (1975: 112), in USNM [# 46867]. Synonymy established by Lindroth (1954b: 122).
Nebria curtulata Casey, 1924: 20. Type locality: «W[est] S[ain]t Modest[e], Labrador» (original citation). Lectotype (♂), designated by Lindroth (1975: 112), in USNM [# 46856]. Synonymy established by Lindroth (1954b: 122).

This subspecies ranges from Greenland (Böcher 1988: 5) to Alaska (Lindroth 1961a: 79), south to northeastern Oregon, northeastern Nevada, western South Dakota, and northeastern New York (Kavanaugh 1978: 731-741).




The name Nebria rufescens (Strøm, 1768) is used by several authors instead of Nebria gyllenhali (Schönherr, 1806). I concur with Kavanaugh (1979a: 111) and Ledoux and Roux (2005: 107) that Carabus rufescens Strøm should be considered a nomen dubium.

Nebria gyllenhali lassenensis Kavanaugh, 1979
Nebria gyllenhali lassenensis Kavanaugh, 1979a: 96. Type locality: «Emerald Lake, Mount Lassen (south slope), Lassen Volcanic National Park [Shasta County], California» (original citation). Holotype (♂) in CAS [# 12511].

This subspecies is restricted to mountains in the southern part of the Cascade Range, south of the lower Columbia River valley, and the northern part of the Sierra Nevada, south to Sonora Pass [see Kavanaugh 1979a: Fig. 54].



Nebria gyllenhali lindrothi Kavanaugh, 1979
Nebria gyllenhali lindrothi Kavanaugh, 1979a: 97. Type locality: «Brooklyn Lake (3, 200 m), Albany County, Wyoming» (original citation). Holotype (♂) in CAS [# 12512].

This subspecies is found in the southern Rocky Mountains from the Medicine Bow Mountains and Sierra Madre of southern Wyoming south to northern New Mexico, west to the Uinta Mountains in eastern Utah and the Chuska Mountains in northeastern Arizona (David H. Kavanaugh pers. comm. 2008) [see Kavanaugh 1979a: Fig. 54].



Nebria nivalis gaspesiana Kavanaugh, 1979
Nebria nivalis gaspesiana Kavanaugh, 1979a: 96. Type locality: «Ruisseau du Diable (980-1, 070 m), Mont Albert, Gaspé-Ouest, Québec» (original citation). Holotype (♂) in CAS [# 12510].

This subspecies is known from western Newfoundland, coastal Labrador (Lindroth 1961a: 82), and the east edge of the Ungava Bay in northern Quebec; isolated on high mountains in the Gaspé Peninsula in Quebec and on Mount Katahdin in Maine [see Kavanaugh 1979a: Fig. 53].



Nebria nivalis nivalis (Paykull, 1790)
Carabus nivalis Paykull, 1790: 52. Type locality: «Lapponiae Lulensis [= Luleå, Norrbotten County, Sweden]» (original citation). Lectotype (♂), designated by Kavanaugh (1979a: 111), in NRSS.
Nebria bifaria Mannerheim, 1853: 120. Type locality: «insula St. Pauli [Alaska]» (original citation). Lectotype (♂), designated by Kavanaugh (1979a: 112), in ZILR. Synonymy established by Lindroth (1961a: 81).
Nebria femoralis Motschulsky, 1859b: 541 [primary homonym of Nebria femoralis Chaudoir, 1843]. Type locality: region of Yakutsk, east-central Siberia, Russia (inferred from title of the paper). Lectotype, designated by Shilenkov (1975: 839), in ZILR. Synonymy established by Jeannel (1937b: 4).
Nebria molbis Motschulsky, 1866: 274. Type locality: «Amérique russe» (original citation). Lectotype, designated by Kavanaugh (1979a: 114), in ZMMU. Synonymy established by Lindroth (1961a: 82).
Nebria femorata Motschulsky, 1866: 275. Type locality: «Sib[eria] bor[eali] Jakutzk [= Yakutsk, Yakutia, Siberia, Russia]» (original citation). Syntype(s) location unknown (possibly in ZILR). Synonymy established by Bänninger (1949: 144). Note. The name Nebria femorata Motschulsky has been interpreted as a replacement name for Nebria femoralis Motschulsky, 1859 by some authors (e.g., Ledoux and Roux 2005: 112). However, there is no indication in Motschulsky (1866) that he proposed the name as a replacement name. The footnote on the same page as the description of femorata indicates that Motschulsky (1866: 275) considered Nebria femoralis Chaudoir as belonging to a different genus, Alpaeus. Moreover in the catalogue of his new genera and species described, Motschulsky (1869: 26) listed his Nebria femoralis and Nebria femorata as different taxa.

This Holarctic subspecies is known from northern Europe to the Bering Sea Coast (Farkač and Janata 2003: 88), and from the Arctic Plains in Alaska to Baffin Island and the western edge of the Ungava Bay in northern Quebec (Kavanaugh 1978: 779-782).


CAN: BC, NT, QC, YT USA: AK – Holarctic


Ledoux and Roux (2005: 112) retained Nebria bifaria Mannerheim, 1853 as a valid subspecies of Nebria nivalis.

[hudsonica group]
Nebria bellorum Kavanaugh, 1979
Nebria lacustris bellorum Kavanaugh, 1979a: 95. Type locality: «West Prong Little Pigeon River (at Chimneys Picnic Area; 3000’) [Sevier County], Great Smoky Mountains National Park, Tennessee» (original citation). Holotype (♂) in CAS [# 12506].

This species is restricted to the Great Smoky Mountains National Park and adjacent mountain ranges in the southern Appalachians [see Kavanaugh 1979a: Fig. 52]. Two specimens labeled from Jefferson County in Colorado and Saint Tammany Parish in Louisiana seen by Kavanaugh (1979a: 96) are likely mislabeled.




This taxon, originally described as a subspecies of Nebria lacustris Casey, has been raised to species status by Kavanaugh et al. (2011).

Nebria gouleti Kavanaugh, 1979
Nebria gouleti Kavanaugh, 1979a: 94. Type locality: «Rattlesnake Creek (3000’), 10 miles s[outh]w[est] of Antone, Asotin County, Washington» (original citation). Holotype (♂) in CAS [# 12504].

This species is restricted to Washington, northern Oregon, and Idaho [see Kavanaugh 1979a: Fig. 51]. Seven specimens labeled from Longview (Highwood River) in southwestern Alberta seen by Kavanaugh (1979a: 95) are listed as doubtful by him; two specimens simply labeled from California are likely mislabeled.



Nebria hudsonica LeConte, 1863
Nebria hudsonica LeConte, 1863c: 3. Type locality: «Saskatchewan, Hudson’s Bay Territory» (original citation), restricted to «North Saskatchewan River at Rocky Mountain House, Alberta» by Kavanaugh (1979a: 113). Lectotype (♂), designated by Kavanaugh (1979a: 112), in MCZ [# 643].

This species ranges from the north shore of Lake Superior in western Ontario to southern Yukon Territory and southeastern Alaska (Skagway, David H. Kavanaugh pers. comm. 2008), south to the Columbia River drainage in northern Oregon and to north-central Utah and central Colorado along the Rocky Mountains (Kavanaugh 1978: 745-753). The records from New Mexico (Snow 1885: 66; Fall and Cockerell 1907: 156) are probably in error.



Nebria lacustris Casey, 1913
Nebria lacustris Casey, 1913: 56. Type locality: «Bayf[ie]ld [Bayfield County], Wis[consin]» (lectotype label). Lectotype (♀), designated by Lindroth (1975: 112), in USNM [# 46865].
Nebria expansa Casey, 1913: 56. Type locality: «Indiana» (original citation for the lectotype), restricted to «Turkey Run State Park, Parke County» by Kavanaugh (1979a: 112). Lectotype (♀), designated by Lindroth (1975: 147), in USNM [# 46864]. Synonymy established by Lindroth (1961a: 77).

This species is found from northern New Brunswick (Restigouche County, CNC) to northern Minnesota, north to southeastern Manitoba, south to east-central Iowa, southern Indiana, and western North Carolina along the Appalachians [see Kavanaugh 1979a: Fig. 52]. The record from “Texas” (Casey 1913: 56) is in error (Kavanaugh 1979a: 112).



Subgenus Nakanebria Ledoux and Roux, 2005
Nakanebria Ledoux and Roux, 2005: 183. Type species: Nebria kurosawai Nakane, 1960 by original designation. Etymology. From the surname of the Japanese coleopterist Takehiko Nakane [1920-1999] and the generic name Nebria [q.v.] [feminine].

Six species in western North America (two species) and the Far East (four species).

Taxonomic Note.

According to Ledoux and Roux (2005: 75), Nakanebria is the sister-group to Reductonebria and the two form the sister-group to Catonebria.

Nebria paradisi Darlington, 1931
Nebria vandykei Darlington, 1930: 104 [primary homonym of Nebria vandykei Bänninger, 1928]. Type locality: «near Paradise Valley (about 6000 feet), Mount Rainier [Pierce County], Washington» (original citation). Holotype (♂) in MCZ [# 35405].
Nebria paradisi Darlington, 1931: 24. Replacement name for Nebria vandykei Darlington, 1930.

This species ranges in the Cascade Range from northwestern Washington to northwestern Oregon [see Kavanaugh 1988: Fig. 19].



Nebria turmaduodecima Kavanaugh, 1981
Nebria turmaduodecima Kavanaugh, 1981b: 436. Type locality: «Caribou Basin (2290 m), Trinity Alps, Siskiyou County, California» (original citation). Holotype (♂) in CAS [# 13729].

This species is endemic to the Trinity Alps in northwestern California [see Kavanaugh 1981b: Fig. 21].



Subgenus Reductonebria Shilenkov, 1975
Reductonebria Shilenkov, 1975: 834. Type species: Nebria ochotica Sahlberg, 1844 by original designation. Etymology. From the Latin reducto (bring back, reduce) and the generic name Nebria [q.v.], possibly alluding to the absence of mid-lateral setae on the pronotum of the species [feminine].

Twenty-eight species in North America (24 species) and Siberia and the Far East (four species).

Faunistic Note.

Nebria carbonaria Eschscholtz was reported from Alaska by Horn (1870b: 104) and LeConte (1878a: 479) and there is a specimen of this species in the LeConte collection labeled “Ins. S. Pauli, ” one of the Pribilof Islands in Alaska. Lindroth (1961a: 74) believes the specimen is probably mislabeled since no other specimens of the species have been found on the island despite extensive search. The species was described from specimens collected “in Kamtschatka, bei St. Peter und Paul” which, according to Lindroth (1961a: 74), refers to the town of Petropavlovsk in Kamtschatka. Until recent specimens are collected on this continent, the species is not listed as a North American resident.

[gregaria group]
Nebria arkansana arkansana Casey, 1913
Nebria arkansana Casey, 1913: 52. Type locality: «Indiana» (original citation for the lectotype), which according to Lindroth (1961a: 70) and Kavanaugh (1979a: 112) is incorrect; «Valley [of the] Upper San Juan [River], [Archuleta County], Color[ado]» selected by Lindroth (1961a: 70). Lectotype (♂), designated by Lindroth (1975: 111), in USNM [# 46858].

This subspecies is found in southern Wyoming, Colorado, northern New Mexico, and southeastern Utah (Kavanaugh 1978: 675-678). One specimen labeled from Nez Perce County in western Idaho is considered doubtful by Kavanaugh (1978: 678).




This species is placed with the species of the lyelli group by Ledoux and Roux (2005: 195).

Nebria arkansana edwardsi Kavanaugh, 1979
Nebria arkansana edwardsi Kavanaugh, 1979a: 100. Type locality: «Logan Pass (7100’), Glacier National Park [Flathead County], Montana» (original citation). Holotype (♂) in CAS [# 12495]. Etymology. The subspecific name was proposed for J. Gordon Edwards [1919-2004], teacher, coleopterist, and mountaineer.

This subspecies is widely distributed in the Rocky Mountain region from southern Yukon Territory south to northeastern Nevada, southern Idaho, and northern Wyoming [see Kavanaugh 1979a: Fig. 58].



Nebria arkansana fragilis Casey, 1924
Nebria fragilis Casey, 1924: 21. Type locality: «North Fork, Provo Cañon [Utah County], Utah» (original citation). Lectotype (♂), designated by Lindroth (1975: 112), in USNM [# 46857].
Nebria arkansana uinta Kavanaugh, 1979a: 102. Type locality: «Lost Lake (9800’), Summit County, Utah» (original citation). Holotype (♂) in CAS [# 12497]. Synonymy established by Kavanaugh (1984: 167).
Nebria fragilis teewinot Kavanaugh, 1979a: 103. Type locality: «Mount Teewinot (southeast slope; 7100-9000’), Grand Teton National Park [Teton County], Wyoming» (original citation). Holotype (♂) in CAS [# 12500]. Synonymy established by Kavanaugh (1984: 167).

This subspecies is restricted to mountains in western Wyoming and northern and central Utah (Kavanaugh 1979a: Figs 58–59).



Nebria arkansana oowah Kavanaugh, 1979
Nebria arkansana oowah Kavanaugh, 1979a: 102. Type locality: «Mill Creek (at Oowah Lake; 8800’), Grand County, Utah» (original citation). Holotype (♂) in CAS [# 12496].

This subspecies is endemic to the La Sal Mountains in eastern Utah [see Kavanaugh 1979a: Fig. 58].



Nebria charlottae Lindroth, 1961
Nebria charlottae Lindroth, 1961a: 67. Type locality: «Queen Charlotte Islands [British Columbia]» (original citation), restricted to «Masset, Graham Island» by Kavanaugh (1992: 55). Holotype (♂) in CNC [# 7611].

This species is restricted to the Queen Charlotte Archipelago (Kavanaugh 1992: 55).