Corresponding author: Yves Bousquet (
Academic editor: Terry Erwin
All scientific names of
One new genus-group taxon,
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
The
The extant hydradephagan families include the
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
This work catalogues all geadephagan taxa of America, north of Mexico. The last catalogue covering the
The first checklist / catalogue covering the North American
In April 1866, John Lawrence LeConte [1825-1883] published the first part of a checklist of the
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
While the
In 1972, Ross H. Arnett, Jr. [1919-1999], the catalyst behind the birth of the Coleopterist’s Society and its journal
In November 1978, the Science and Educational Administration, USDA, released its first fascicle, covering the family
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
The first catalogue of the world
A comparison of the number of valid species and genera between this and previous checklists / catalogues is presented in
North American
|
|
|
|
|
|
---|---|---|---|---|---|
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 |
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)
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
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 (
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.
Collections cited in the catalogue are referred to by the abbreviations listed below.
Besides those used for provinces and states (see
In addition, the International Commission on Zoological Nomenclature is sometimes abridged to “Commission” and United States of America to “United States.”
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
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
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.
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.
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
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
Harris, well known for his work in economic entomology (his profile having appeared on every cover of the
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).
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
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
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.
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).
Although Ulke described only two North American carabids in his life,
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
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 death
LeConte used small colored paper disks to indicate the provenance of his specimens. The color system used is as follows:
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
From 1884 to the end of his life, Casey described 1,864 new species-group taxa of North American
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).
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.
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.
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
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
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.
Unfortunately, there is no consensus among coleopterists concerning the classification of
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
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
The
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,
Traditionally ranked as a distinct family, rhysodids (also known as wrinkled bark beetles) have been included within the family
About 355 species of rhysodids are currently known and are placed into seven family-group taxa, namely
Tribe
Tribe
Tribe
Tribe
Tribe
Tribe
Tribe
Monophyly of the
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
Fossils belonging to the family
The world classification of family-group taxa, which has been adopted for the North American fauna in this catalogue, is outlined in
Classification of world family-group taxa of
Subfamily |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Tribe |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Subfamily |
|
Tribe |
|
Tribe |
|
Subfamily |
|
Supertribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Supertribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
|
Tribe |
This subfamily includes the tribes
The subfamilies
Tribe
Tribe
Tribe
Tribe
Tribe
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.
This subfamily includes two species,
This subfamily contains about 1,300 species (Lorenz 2005: [i]) placed in four tribes:
Tribe
Relationships among the four genera have not been investigated. Whether or not
Tribe
Tribe
Tribe
The supraspecific classification of
Based on morphological (larvae and endophallus of adults) and molecular sequence data, Deuve (2004) recognized eight major lineages within the genus
Jeannel (1940: 68) recognized two major lineages within the calosomatines: the
The systematic position of
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
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,
Tribe
Tribe
Tribe
Tribe
Tribe
Tribe
This subfamily contains a single genus,
Recently Sciaky and
Klausnitzer (2003) described a new species,
This subfamily includes a single tribe with obscure relationships. Bell (1967) listed the
The tribe is represented only in the Northern Hemisphere and includes three genera, all represented in the Nearctic Region.
This subfamily includes a single genus,
The genus
This subfamily currently consists of two tribes:
Tribe
Tribe
This subfamily includes two genera,
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:
Tribe
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
Tribe
Tribe
The North American fauna is represented by seven species of
Tribe
At least three putative clades are recognized within the tribe and are usually ranked as subtribes. The forcipatorines, exclusively Neotropical, include the genera
Iablokoff-Khnzorian (1960: 93) described a new genus,
Tribe
Tribe
Tribe
McKay (1991) described a fossil from Cretaceous crater lake deposits at Orapa, Botswana, under the name
Tribe
This subfamily includes a single tribe with about 290 species in 34 genera, arrayed in five subtribes (see Roig-Juñent 2000):
Roig-Juñent’s (2000) parsimony analysis based on morphological characters of adults suggested that the three native North American genera (
This subfamily includes a single genus,
This subfamily includes three genera, each arrayed in its own tribe:
Tribe
Tribe
Tribe
This subfamily includes one tribe with two genera:
Relationships of the subfamily are unclear. Liebherr and Will (1998: 150) suggested that
This group has been reported in publications of the xix and early xx Centuries under the name
This subfamily includes three genera placed in two subtribes:
Several authors agree that the tribes
As discussed under
Tribe
Based on larval character states, Grebennikov and Maddison (2005) suggested that
The North American fauna comprises about 225 species grouped into nine genera, all belonging to the subtribe
Tribe
Tribe
Tribe
This subfamily, which includes the tribes
This subfamily is found in the Northern Hemisphere and Oriental Region. Only the tribe
Tribe
Tribe
Following Maddison and Ober (2011: 237), this subfamily is restricted to the tribe
Members of this subfamily were traditionally included in the
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):
The genus
Tribe
Tribe
This subfamily includes a single species,
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 {
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
Tribe
Tribe
Tribe
Tribe
Tribe
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:
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
Tribe
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
Unlike most carabid larvae, those of brachinines are ectoparasites and feed on carabid and water beetle pupae.
Harpalinae is the largest subfamily of
In this catalogue, members of
Tribe
Moore (1965: 5) included the Australian genus
Tribe
Tribe
Tribe
Alluaud (1935: 28) reported that one of his colleagues rearing pupae of various insect groups for parasitic
Tribe
Tribe
Tribe
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
Tribe
Tribe
This tribe has been known in the past under the name
Tribe
The name
Tribe
Tribe
Jeannel (1948a: 627) indicated that the genus
Tribe
Straneo (1950: 65) first included the genus
Tribe
Some groups, such as dercylines, melanchitonines, and geobaenines, are sometimes included within the
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 (
Tribe
Peleciines are restricted to the Southern Hemisphere. Straneo and Ball (1989) recognized two subtribes:
Tribe
Tribe
Tribe
Jeannel (1949a: 849) associated the genus
Tribe
Two major groups among
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
Tribe
Tribe
Tribe
Tribe
Tribe
Tribe
The 235 or so species are arrayed in 23 genera distributed among four subtribes following Ball (1992a).
Tribe
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
Tribe
Tribe
Tribe
Tribe
Relationships among the North American genus-group taxa have been addressed but are still inadequately understood. According to Liebherr (1991b: 5),
Tribe
The North American fauna includes two species of the genus
Tribe
Tribe
Tribe
Tribe
Tribe
The
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
Tribe
This tribe includes close to 170 species in the world. Only six, all in the genus
Tribe
The tribe is represented in North America by six species belonging to the New World genus
Tribe
Tribe
The main characteristic of the
Tribe
Tribe
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 (
Tribe
Tribe
Tribe
Tribe
Tribe
Tribe
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):
Tribe
Tribe
Tribe
The hypogean and monospecific genus
Tribe
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
Tribe
Tribe
Sloane (1914) and Reichardt (1974) recognized two subtribes among helluonines:
Tribe
Tribe
The North American fauna currently consists of 2,676 valid species-group taxa (2,439 species) of
Among the 62 species accidentally introduced,
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 (
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).
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.
List of species-group taxa adventive and established in North America.
|
|
|
||
---|---|---|---|---|
|
|
|
||
|
||||
• | 1977 | |||
• | 2007 | |||
|
||||
• | • | 1923 | ||
• | 1968 | |||
|
||||
• | N/A | |||
• | • | 1890 | ||
• | • | 1890 | ||
• | N/A | |||
|
||||
• | 1948 | |||
• | • | <1838 | ||
• | • | 1915 | ||
|
||||
• | 1978 | |||
|
||||
• | 1987 | |||
|
||||
• | 1933 | |||
• | 1927 | |||
• | 1965 | |||
• | <1863 | |||
|
||||
• | 1930 | |||
• | • | 1947 | ||
• | 1942 | |||
• | 1956 | |||
• | <1897 | |||
• | 1891 | |||
• | 1907 | |||
• | 1967 | |||
• | • | <1823 | ||
• | 1940 | |||
• | • | <1900 | ||
|
||||
• | 1984 | |||
• | 1997 | |||
• | • | 1937 | ||
• | • | 1926 | ||
• | 1965 | |||
|
||||
• | 1929 | |||
• | ? | <1865 | ||
• | 1905 | |||
• | 1929 | |||
• | • | 1925 | ||
• | • | 1904 | ||
• | • | 1929 | ||
• | 1988 | |||
• | 1971 | |||
• | • | 1901 | ||
|
||||
• | 1911 | |||
• | 1951 | |||
• | • | 1931 | ||
• | 1954 | |||
• | 1953 | |||
• | 1937 | |||
• | • | <1798 | ||
• | 1981 | |||
|
||||
• | 1928 | |||
• | <1874 | |||
• | ? | <1894 | ||
• | <1835 | |||
• | • | 1840 | ||
• | 1931 | |||
|
||||
• | • | <1853 | ||
|
||||
• | 2007 | |||
|
||||
• | 1992 | |||
• | ? | |||
• | 1952 | |||
• | 1996 | |||
• | ? | • | <1833 |
YFC: Year of first collection
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:
Number of adventive species of
Number of North American species-group taxa of
Cumulative number of species-group taxa of
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.
Species-group taxon statistics for political regions.
|
|
|
|
|
---|---|---|---|---|
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
Checklists and taxonomic treatments on
|
|
---|---|
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) |
Six species in western North America (three species), South America (two species), and northern Eurasia (one species). The species are arrayed in two genera:
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.
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.
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).
This species is known only from the western regions of Washington and Oregon (Lindroth 1961a: 4).
Worldwide, with about 355 species arrayed in six tribes:
Bell (1970) revised the North American, Middle American, and West Indies species and provided keys for their identification.
Worldwide, with about 135 species arrayed in three genera:
About 75 species in the Nearctic (six species), Neotropical (about 65 species), and Palaearctic (three species) Regions arrayed in four subgenera:
Bell and Bell (1985) revised the species of the world and provided keys for their identification.
Northern Hemisphere, with nine species in North America (six species), Japan (one species), Caucasian region (one species), and southern Europe (one species).
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.
This subspecies is known only from the type locality in northern Georgia (Bell and Bell 1985: 90).
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.
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).
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).
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.
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).
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.
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.
Three species in North American (two species) and Japan (one species).
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).
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.
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:
This tribe contains a single genus.
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.
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).
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).
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:
Bousquet and Smetana (1996) reviewed the species and provided a key for their identification.
One North American species.
The species was treated in Lindroth’s (1961a: 88-90) monograph on the
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).
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:
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:
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
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.
Three western North American species.
Erwin (1970b) revised the species and provided a key for their identification.
Perrault (1991a) added three species from the Far East (
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).
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).
This species is known only from a few localities along the Pacific Coast in northern California [see Erwin 1970b: Fig. 7].
Seven species in western North America (three species), Japan (three species), and Jilin Province in China (one species) arrayed in two subgenera:
Kavanaugh (1995, 1996) regarded
Three species in western North America.
Ledoux and Roux (2005: 712) reviewed the species and provided a key for their identification.
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].
This species ranges from the Three Brothers Mountain in southern British Columbia south to Mount Baker in Washington [see Kavanaugh 1984: Fig. 31].
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).
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
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).
The species of
Thirty-one species (Ledoux and Roux 2005: 82) in North America (seven species) and Eurasia (26 species). Two species are Holarctic (
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].
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].
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).
This species is placed in its own group by Ledoux and Roux (2005: 82).
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
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].
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].
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].
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).
Ledoux and Roux (2005: 112) retained
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
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.
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.
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).
Six species in western North America (two species) and the Far East (four species).
According to Ledoux and Roux (2005: 75),
This species ranges in the Cascade Range from northwestern Washington to northwestern Oregon [see Kavanaugh 1988: Fig. 19].
This species is endemic to the Trinity Alps in northwestern California [see Kavanaugh 1981b: Fig. 21].
Twenty-eight species in North America (24 species) and Siberia and the Far East (four species).
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
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].
This subspecies is restricted to mountains in western Wyoming and northern and central Utah (Kavanaugh 1979a: Figs 58–59).
This subspecies is endemic to the La Sal Mountains in eastern Utah [see Kavanaugh 1979a: Fig. 58].
This species is restricted to the Queen Charlotte Archipelago (Kavanaugh 1992: 55).
This species is endemic to the Aleutian Islands [see Kavanaugh 1981a: Fig. 8].
This species is endemic to high elevations in the Queen Charlotte Islands [see Kavanaugh 1984: Fig. 31] and adjacent mainland on Mount McNeil (Kavanaugh 1992: 56) in British Columbia.
Clarke et al. (2001: 1416) concluded that this taxon may be more appropriately treated as a subspecies of
This species is known only from the Steens Mountains in south-central Oregon [see Kavanaugh 1984: Fig. 31].
This species is placed with the species of the
This species is known only from a small area in the Alexander Archipelago and northwestern British Columbia [see Kavanaugh 1988: Fig. 24].
This species is known from several islands in the Queen Charlotte Archipelago, British Columbia (Kavanaugh 1992: 55).
Based on DNA sequence analyses, Clarke et al. (2001: 1416) concluded that this taxon may not represent a distinct taxonomic unit but rather a variant of
This subspecies is known only from the Warner Mountains in northeastern California [see Kavanaugh 1979a: Fig. 56].
This subspecies ranges from the Aleutian Islands in Alaska (Lindroth 1961a: 70) to southwestern Northwest Territories (Tungsten, David H. Kavanaugh pers. comm. 2008), south to northwestern Montana (Edwards 1975: 50) and southern Oregon [see Kavanaugh 1988: Fig. 20]. The records from New Mexico (Snow 1885: 66; Fall and Cockerell 1907: 156), Colorado (Wickham 1902: 232; Armin 1963: 94), and Wyoming (Lavigne 1977: 46) are probably in error.
This species is placed in the
This subspecies is yet recorded only from the Klamath Mountains system in northwestern California [see Kavanaugh 1979a: Fig. 56].
This subspecies is restricted to the Pine Valley Mountains in southwestern Utah [see Kavanaugh 1979a: Fig. 60].
This subspecies is known only from mountains in southwestern Utah [see Kavanaugh 1979a: Fig. 60].
This species is placed in the
This subspecies ranges from the Kenai Peninsula in Alaska (Kavanaugh 1978: 671) and southwestern Yukon Territory (Kluane Lake, Sydney G. Cannings pers. comm. 2009) south to the southern part of the Cascade Range and the Sierra Nevada in central California (Kavanaugh 1978: 671).
This subspecies is endemic to the Olympic Peninsula in Washington [see Kavanaugh 1979a: Fig. 55].
As far as known, this subspecies is restricted to the Sierra Nevada, between Sonora Pass and Buckeye Pass, in California [see Kavanaugh 1981b: Fig. 21].
This species is endemic to high elevations in the Olympic Mountains in northwestern Washington [see Kavanaugh 1984: Fig. 21].
This species is known only from Mount Lyell at the edge of the Yosemite National Park in the Sierra Nevada, California (Kavanaugh 1978: 766).
This species is known only from the Wallowa Mountains in northeastern Oregon [see Kavanaugh 1984: Fig. 31].
This species is known only from the canyon of the South Fork of the American River in the Sierra Nevada, California [see Kavanaugh 1979a: Fig. 62].
This species is known only from the type locality in south-central Utah.
This species ranges along the Pacific Coast from the Queen Charlotte Islands to northern California (Kavanaugh 1992: 57).
This species is placed in a separate group (
The range of this species extends from northwestern Washington to northeastern Idaho, south to southern California [see Kavanaugh 1979b: Fig. 2]. The records from Colorado (Wickham 1902: 232; Armin 1963: 89) and Montana (Hatch 1933a: 7) must be in error.
This species is known only from the original two specimens collected 141 kilometers apart in the Grand Canyon National Park (Kavanaugh 2008: 5).
This species ranges from the Kenai Peninsula in southern Alaska (Lindroth 1961a: 75) south to western Idaho and central Oregon (Kavanaugh 1978: 766-772). According to Kavanaugh (1978: 772), a number of specimens labeled from “California,” Nevada, and Yukon Territory are doubtful. The records from the Absaroka Range in south-central Montana (Hatch 1933a: 7) and western San Juan Mountains in Colorado (Wickham 1902: 232) are also doubtful.
This species is known only from the vicinity of the type locality in northeastern Arizona [see Kavanaugh 1979a: Fig. 63].
This species is known from the southern Appalachian Mountains in western North Carolina and eastern Tennessee (Kavanaugh 1978: 674).
This species is placed in a distinct group (
This subspecies, as far as known, is endemic to the Chuska Mountains in northeastern Arizona [see Kavanaugh 1979a: Fig. 61].
This subspecies ranges from the west edge of the Wrangell-St. Elias National Park in southeastern Alaska to the Great Slake Lake in Northwest Territories, south to northern New Mexico, northern Arizona, central Nevada, and the lower eastern slope of the Sierra Nevada in California, east to western South Dakota and western Nebraska [see Kavanaugh 1979b: Fig. 4].
This species is found east of the Mississippi River from Nova Scotia (Lindroth 1961a: 77) to north-central Illinois (Kavanaugh 1978: 800), south to northern Alabama (Löding 1945: 12), northeastern Georgia (Leng 1910: 73; Fattig 1949: 12), and east-central South Carolina (Ciegler 2000: 20) along the Appalachian Mountains. The record from east-central Missouri (Summers 1873: 133) needs confirmation; that from Colorado (Elias 1987: 632) is likely based on a mislabeled specimen; that from northern Wisconsin along Lake Superior (Wickham 1896c: 131) probably refers to
This species is placed in a special group (
This species ranges from the coast of Labrador and the Ungava Bay region in northern Quebec to the Rocky Mountains in western Alberta, south to central Colorado, the Adirondack Mountains in northeastern New York, and New England [see Kavanaugh 1979b: Fig. 3].
Thirty-one species (Ledoux and Roux 2005: 240) in North America (20 species) and Siberia and the Far East (11 species).
As far as known, this species is endemic to the Wallowa Mountains in northeastern Oregon [see Kavanaugh 1984: Fig. 31].
This subspecies is known only from the western slope of the White Mountains in eastern California [see Kavanaugh 1984: Fig. 21].
This subspecies has been found only on the eastern slope of the southern Sierra Nevada in California [see Kavanaugh 1979a: Fig. 67].
This subspecies is known from the Skagway area in southeastern Alaska and northwestern British Columbia south along the Cascade Range to Mount Shasta in north-central California [see Kavanaugh 1979a: Fig. 67]. The record from “Whitehorse Pass,” Yukon Territory (Kavanaugh 1978: 773), refers to the Skagway area in Alaska (Sydney G. Cannings pers. comm. 2009).
This subspecies in known from Vancouver Island and the Olympic Peninsula in northwestern Washington [see Kavanaugh 1979a: Fig. 67].
This species ranges from the Aleutian Islands south to western Montana (Hatch 1939a: 118) and southern Washington (Kavanaugh 1978: 775-778). A few specimens labeled from Arizona, California, and Oregon are known (Kavanaugh 1978: 778) but considered doubtful. The record from Colorado (Elias 1987: 632) is in error (David H. Kavanaugh pers. comm. 2012).
This species is found in the mountains of south-central and western Idaho [see Kavanaugh 1979a: Fig. 65] from the Seven Devils Mountains in the north to the Sawtooth Range in the south.
This subspecies is known only from the type locality in the White Mountains of eastern California.
This subspecies is found from southern British Columbia, including southern Vancouver Island, south along the Cascade Range to central Oregon [see Kavanaugh 1979a: Fig. 64]. One specimen labeled from Leavenworth Valley in Colorado seen by Kavanaugh (1979a: 106) is likely mislabeled.
This subspecies is yet recorded only from the type locality in the Strawberry Mountains of eastern Oregon [see Kavanaugh 1979a: Fig. 64].
This subspecies occurs along the Cordilleras from the Alexander Archipelago to southernmost Yukon Territory (Lindroth 1961a: 84), south to southwestern Montana, south-central Idaho, and northeastern Oregon [see Kavanaugh 1979a: Fig. 64]. The record from northern Colorado (Armin 1963: 89) is probably in error.
This species is placed in its own group by Ledoux and Roux (2005: 240).
This subspecies is endemic to the Sierra Nevada and adjacent mountains in western Nevada [see Kavanaugh 1979a: Fig. 64].
This subspecies is restricted to the Klamath Mountains system of the Coast Ranges in southwestern Oregon and northwestern California [see Kavanaugh 1979a: Fig. 64].
This subspecies is restricted to the Cascade Range of central Washington and northern Oregon [see Kavanaugh 1979a: Fig. 65].
This subspecies ranges along the mountains of the Pacific Coast from the Alexander Archipelago to northern Washington [see Kavanaugh 1979a: Fig. 65].
This species is known from the Sierra Nevada in California (Erwin and Ball 1972: 81) and adjacent mountains in Washoe County, western Nevada (Kavanaugh 1978: 798).
This subspecies is restricted to the main chain and Eastern Divide of the Sierra Nevada of California [see Kavanaugh 1979a: Fig. 66; David H. Kavanaugh pers. comm. 2012].
This subspecies is endemic to the Western Divide of the southern Sierra Nevada of California [see Kavanaugh 1979a: Fig. 66; David H. Kavanaugh pers. comm. 2012].
This species is yet known only from the type locality in northeastern Arizona [see Kavanaugh 1984: Fig. 32].
This species is restricted to the San Juan Mountains in southwestern Colorado and the Abajo Mountains in southeastern Utah (Erwin and Ball 1972: 97).
This species is found in the Rocky Mountains in southern Wyoming and Colorado (Erwin and Ball 1972: 97).
This taxon has been considered a synonym of
This subspecies is restricted to the Sierra Nevada in Fresno, Inyo, and Tulare Counties, California (Kavanaugh 1978: 753-754).
This subspecies is known only from Mono and Tuolumne Counties in the Sierra Nevada of California (Kavanaugh 1978: 754; David H. Kavanaugh pers. comm. 2012).
This species ranges along the mountains paralleling the Pacific Coast from southeastern Alaska along the British Columbia border (Lindroth 1961a: 86) and Kluane National Park in southwestern Yukon Territory (David H. Kavanaugh pers. comm. 2010) to west-central Oregon (Kavanaugh 1978: 811).
This species is placed in the
This subspecies is known only from Beaver and Piute Counties in southern Utah (Kavanaugh 1978: 813).
This subspecies is known only from southern Utah [see Kavanaugh 1984: Fig. 32]. One specimen labeled from Navajo County in Arizona seen by Kavanaugh (1984: 165) is likely mislabeled.
This subspecies is restricted to the Henry Mountains in south-central Utah [see Kavanaugh 1979a: Fig. 70].
This species is endemic to the Trinity Alps of northwestern California.
This species is found along the Rocky Mountains in Colorado and New Mexico (Kavanaugh 1978: 813-815). The record from Idaho (LeConte 1879d: 500) is probably in error.
This beautiful subspecies is recorded only from the type locality in Gros Ventre Mountains, western Wyoming [see Kavanaugh 1979a: Fig. 69].
This subspecies is restricted to the Rocky Mountains in western Alberta and southeastern British Columbia [see Kavanaugh 1979a: Fig. 69].
This species is placed in the
This species is known only from the Sierra Blanca and Capitan Mountains in central New Mexico [see Kavanaugh 1984: Fig. 32].
This species is endemic to the Steens Mountains in south-central Oregon [see Kavanaugh 1984: Fig. 32].
This subspecies ranges from Medicine Bow and Sierra Madre Ranges in southern Wyoming south to the Rampart Range in central Colorado [see Kavanaugh 1984: Fig. 32].
This subspecies ranges along the Rocky Mountains from southern Montana and southeastern Idaho south to northeastern Nevada, southern Utah, and west-central Wyoming [see Erwin and Ball 1972: Fig. 42)].
The form
This subspecies is known only from the Olympic Mountains and the Cascade Range of Washington [see Kavanaugh and Schoville 2009: Fig. 13].
This subspecies is restricted to the Cascade Range in Oregon from Mount Hood south to the Three Sisters area [see Kavanaugh and Schoville 2009: Fig. 13].
Sixty-five species in the Palaearctic Region, one of them adventive in North America.
This Palaearctic species is adventive in North America where it is known from the Willamette Valley in northwestern Oregon and southwestern Washington [see LaBonte 2011: Fig. 4]. Since over 3,000 specimens have been collected, the species is clearly established in the region. The first inventoried specimen found in the area was caught in late 2007 (Kavanaugh and LaBonte 2008: 482). The species has also been collected twice in eastern North America, in Quebec in 1930 and Saint Pierre and Miquelon in 1937 (Lindroth 1961a: 78) but it did not become established in this part of the continent.
This tribe includes a single genus.
About 55 species in the arctic, subarctic, boreal, and temperate areas of the Nearctic (15 species, of which two are adventive), Neotropical (two species in mountains of Middle America), and Palaearctic (about 40 species) Regions, including northern Africa (see Barševskis 2007). Two species,
Lindroth (1961a) reviewed the North American species and provided a key for their identification. One adventive species (
This species ranges from Nova Scotia (Lindroth 1961a: 94) to southeastern Minnesota (Donald P. Schwert pers. comm. 1989), south at least to southeastern Nebraska (Richardson County, Foster F. Purrington pers. comm. 2009), central Missouri (Boone County, CMNH), northeastern Georgia (Leng 1910: 73; Fattig 1949: 11), and southwestern South Carolina (Ciegler 2000: 20). Two specimens labeled from the Santa Catalina Mountains in Arizona (MCZ, collection Fall) are known and so the species probably ranges farther west in southern United States.
The range of this circumpolar species extends from Iceland to the Bering Sea Coast (Bousquet and Barševskis 2003: 96) and from Alaska (Lindroth 1961a: 95) to Newfoundland (Lindroth 1955a: 34), south to Pennsylvania (Capogreco 1989b: 4) and to New Mexico and Arizona (Fall 1906: 85; Lindroth 1961a: 95) along the Rocky Mountains. Fossil remnants from a Plio-Pleistocene sequence have been found in northwestern Greenland (Böcher 1995: 18).
This Palaearctic species is adventive in North America where it is known from Newfoundland and Saint Pierre and Miquelon (Lindroth 1955a: 37) to the Gaspé Peninsula in Quebec (LeSage 1996: 23), south to Connecticut (Krinsky and Oliver 2004: 396) and “Rhode Island” (Sikes 2003: 7), and from southwestern British Columbia (Lindroth 1961a: 101). The first inventoried specimen found on the east side of this continent was caught in Newfoundland in 1923 (Brown 1950b: 197) and on the west coast in Vancouver in 1957 (Lindroth 1961a: 101). Dejean’s (1831: 589) record of this species from «Amérique septentrionale» almost certainly refers to
This species is known from the Far East (Bousquet and Barševskis 2003: 96) and from Alaska (Lindroth 1961a: 97) to Newfoundland (Lindroth 1955a: 36); isolated on some mountains of New England (Lindroth 1961a: 96-97) and New York (Essex County, CNC, MCZ), and also known from northern Wyoming (Johnson County, CMNH). Fossil remnants of this species, dated between about 16,700 and 18,100 years B.P., have been unearthed in southeastern Iowa (Baker et al. 1986: 96).
This species occurs in the western mountain ranges, from southwestern Alberta and southern British Columbia, south to northeastern California (Lindroth 1961a: 98), northeastern Arizona, and northern New Mexico [see Morgan and Morgan 1979: Fig. 5]. Fossil remnants, dated about 11,800 years B.P., have been unearthed in northeastern Wisconsin (Morgan and Morgan 1979: 232).
This species is known from scattered localities from Newfoundland (Lindroth 1955a: 37) to the Gulf of Alaska coast (Lindroth 1961a: 98); also known from northwestern Minnesota (Clearwater County, CNC).
This species is restricted to some mountains in New England (Lindroth 1961a: 100) and the Adirondacks in northeastern New York (Notman 1928: 211).
This species is found from Vancouver Island (Lindroth 1961a: 99) to northwestern Montana (Russell 1968: 46), south to southern Oregon (Harney County, CMNH). The record from Texas (Fall 1906: 90) is probably in error.
The range of this species extends from western Maine (Oxford County, André Larochelle pers. comm. 1990) to “South Dakota” (Kirk and Balsbaugh 1975: 15), south to east-central Texas (Riley 2011) and northern Florida (Peck and Thomas 1998: 15), west along the southwest to “Arizona” (Lindroth 1961a: 101) including northwestern Colorado (Barševskis 2009: 138). The record from Cape Breton Island in Nova Scotia (McCorquodale 2000: 339) is based on a misidentified specimen of
This Palaearctic species is adventive in North America where it is known only from Nova Scotia, including Cape Breton Island (McCorquodale 2000: 339, as
This species is known from California (Fall 1906: 91; Lindroth 1961a: 99), as far north as the San Francisco Bay area (Alameda County, MCZ), Arizona (Wickham 1898: 300; Lindroth 1961a: 99), and the states of Sonora and Chihuahua in Mexico (Erwin 2007a: 59). The record from “Oregon” (Fall 1906: 91) needs confirmation.
This species is found from Labrador to central Alaska, south to northern British Columbia (Lindroth 1961a: 94-95), New Mexico (Fall 1906: 84; Lindroth 1961a: 94) along the Rocky Mountains, Oklahoma (Latimer County, UASM), southwestern Arkansas (Columbia County, INHS), east-central Alabama (Lee County, CNC), central Georgia (Fattig 1949: 11), and southern South Carolina (Ciegler 2000: 20); also recorded from Sakhalin Island (Barševskis 2001: 38).
As far as known, this species is restricted to the Sierra Nevada in California (Fall 1906: 90; Lindroth 1961a: 99).
This species is found in the western mountain ranges from western Alberta and British Columbia (Lindroth 1961a: 97) south at least to central Oregon (Crook County, CNC) and southwestern Colorado (Elias 1987: 632). The record from Houston, Texas (Fall 1906: 87) needs confirmation; that from Alaska (Fall 1926a: 129) refers to
This species ranges from southern Alaska (Lindroth 1961a: 100) to northern California, at least as far south as Mendocino County (Fall 1906: 89), east to northern Idaho (Hatch 1953: 61). Fossil remnants, dated from the Late Pleistocene, have been unearthed in northeastern Siberia (Kiselev 1981: 12).
Worldwide, with about 1,300 species (Lorenz 2005: [i]) arrayed in four tribes:
Northern Hemisphere and mountains in Mexico, with about 200 species arrayed by most authors in four genera:
Six species restricted to the boreal and temperate regions of eastern North America.
Lindroth’s (1961a) key covers four of the species currently recognized;
This species is found along the southern parts of the Appalachians from North Carolina and Tennessee (Darlington 1932: 152) to northern Alabama (Löding 1945: 11) and northeastern Georgia (LeConte 1853c: 399; Leng 1910: 73).
Roeschke (1907a: 263) and Lindroth (1961a: 29) regarded this form as a subspecies of
The range of this subspecies extends from Cape Breton Island to southern Ontario (Lindroth 1961a: 28), south to the Black Mountains in North Carolina (Darlington 1933a: 64). The record from “South Carolina” (Bousquet and Larochelle 1993: 78) refers to
This subspecies occurs in the mountains south of the French Broad River (Barr 1969: 76) from southwestern North Carolina and adjacent parts of Tennessee (Darlington 1933a: 64) to northeastern Georgia (Fattig 1949: 9) and northwestern South Carolina (Ciegler 2000: 30).
This species ranges from northwestern Ohio (Barr 1974a) to central Indiana (Blatchley 1910: 42), south to north-central Mississippi (Grenada County, Drew A. Hildebrandt pers. comm. 2009) and southwestern Virginia (Barr 1974a).
This species ranges from Newfoundland (Lindroth 1955a: 23) to east-central Saskatchewan (Hooper 1978: 19), south to northeastern Minnesota (Gandhi et al. 2005: 923), northern Wisconsin along Lake Superior (Wickham 1896c: 131), the Adirondack Mountains in northeastern New York (Notman 1928: 208), and New England [see Lindroth 1963a: Fig. 68]. The record from southwestern Ohio (Dury 1906: 257) is probably in error. Roeschke (1907a: 268) statement that this species is found “Küstengebiet von Maine bis Washington D.C.” is in error. Fossil remnants, dated between 22,200 and 25,200 years B.P., have been unearthed in central Illinois (Schwert 1992: 76).
Lindroth (1961a: 29) retained
This species is known from a small area from Michigan to northern Virginia (Barr 1974a) and western Maryland (Bailey et al. 1994: 320). The record from “Illinois” (Bousquet and Larochelle 1993: 78) is probably in error.
Roeschke (1907a: 266), Lindroth (1961a: 29), Lorenz (2005: 64), and Erwin (2007a: 170) listed this taxon as a subspecies of
This subspecies is found from Newfoundland (Lindroth 1955a: 21) to southeastern Manitoba (Lindroth 1961a: 29), south to east-central Iowa (Wickham 1888: 81, as
This mainly coastal subspecies ranges from southeastern New York (Barr 1974a) and southeastern Pennsylvania (York County, CMNH) south to Wake County in east-central North Carolina (Brimley 1938: 115). The record from northern Georgia (Fattig 1949: 9) probably refers to
Northern Hemisphere, with about 115 species (Lorenz 2005: 64-66) in the Nearctic (two western species) and Palaearctic Regions.
Gidaspow (1973) revised the North American species. Lindroth (1961a: 25-26) covered the species in his monograph of the Canadian and Alaskan
This subspecies is known from southwestern Idaho, western Wyoming, and northern Utah [see Gidaspow 1973: Fig. 7].
This subspecies is found from southeastern British Columbia south to northeastern Oregon (LaBonte 1988: 264), northern Idaho, and the Bitter Root Mountains in western Montana (Roeschke 1907a: 227) [see Gidaspow 1973: Fig. 7]. One specimen labeled from Logan Canyon in northern Utah is also known (Gidaspow 1973: 98).
Lindroth (1961a: 26) treated this form as a valid species but both Roeschke (1907a: 227) and Gidaspow (1973: 96) regarded it as a subspecies of
This species ranges from British Columbia, including the Queen Charlotte Islands and Vancouver Island, south at least to southern Oregon [see Gidaspow 1973: Fig. 7], possibly even Mendocino County in northern California (Roeschke 1907a: 229). The record from “Alaska” (Roeschke 1907a: 229) is likely in error.
Fifty-five species in the boreal and temperate regions of North America and the Sierra Madre Occidental in Mexico.
Roeschke (1907a) reviewed the species known at the time.
Nine species in North America (seven species) and the Sierra Madre Occidental of Mexico (two species).
Van Dyke (1938) reviewed the species and provided a key for their identification. One new species was described subsequently by Allen and Carlton in 1988.
The range of this subspecies extends from southern Manitoba south to “Iowa” (Lindroth 1961a: 18; Esau and Peters 1975: 510), northwestern Nebraska, and north-central New Mexico (Van Dyke 1938: 123-124). The record from Wisconsin (Rauterberg 1885: 12, as
This subspecies ranges from southern New Hampshire (Merrimack and Strafford Counties, Donald S. Chandler pers. comm. 2008) south to the Florida Panhandle (Peck and Thomas 1998: 16), west to eastern Texas (Lamar and Fort Bend Counties, Brian Raber pers. comm. 2010; Riley 2011). The records from Maine (Bousquet and Larochelle 1993: 79) and Lawrence in Kansas (Van Dyke 1938: 120) need confirmation.
This subspecies ranges from Nebraska to Ohio, south to Tennessee, Arkansas (Van Dyke 1938: 121), and southwestern Oklahoma (Kondratieff et al. 2005: 173). The record from “Louisiana” (Roeschke 1907a: 142) needs confirmation.
Van Dyke (1938: 121) noted that “the subspecies grades gradually into typical
This subspecies is known so far only from the holotype collected in southeastern Virginia.
This subspecies is known so far only from the holotype and several specimens collected at the type locality (Eric van den Berghe and Robert L. Davidson pers. comm.) in southern New Mexico.
According to Roeschke (1907a: 141), this subspecies is found along the coast from New Jersey to South Carolina. Van Dyke (1938: 122) saw specimens from Virginia and North Carolina and noted that “it is apparently confined to the more maritime or coastal portion of the South Atlantic states, replacing almost entirely the [nomino]typical phase in its area of distribution.”
This species is known only from mountains in southwestern New Mexico in Grant County and other places in the neighborhood of Silver City (Van Dyke 1938: 104).
This subspecies is confined to the Rincon and Huachuca Mountains of southern Arizona (Ball 1966c: 711-712).
This subspecies is known from the Santa Catalina Mountains in Pima County (Ball 1966c: 711) and Aravaipa Canyon in Pinal County (McCleve 1979: 452), southern Arizona.
This subspecies is confined to the Chiricahua Mountains in Cochise County, southeastern Arizona (Ball 1966c: 714).
This subspecies is found in the Pinaleño and White Mountains of southeastern Arizona (Ball 1966c: 713-714).
This subspecies is known only from Mount Wrightson in the Santa Rita Mountains, southern Arizona (Ball 1966c: 715).
This subspecies is confined to the Pinal and Sierra Ancha Mountains in Gila County, eastern Arizona (Ball 1966c: 709).
This subspecies is restricted to high mountains in Coconino County, northern Arizona (Van Dyke 1938: 103).
Van Dyke (1938: 102) treated this form as a species but Ball (1966c: 693) retained it as a subspecies of
This subspecies is found in western New Mexico, eastern Arizona, southwestern Colorado, and southeastern Utah (Van Dyke 1938: 100).
This species is confined to southern Coconino, Yavapai, and Gila Counties in Arizona (Van Dyke 1938: 106).
Van Dyke (1938: 106) retained
This species is known only from the type locality in western Arkansas.
This species ranges from New Jersey (Smith 1910: 199) and Maryland (Van Dyke 1938: 128) to southwestern Illinois (Jackson County, Robert L. Davidson pers. comm. 2012), south to east-central Louisiana (West Feliciana Parish, Igor M. Sokolov pers. comm. 2009), northeastern Mississippi (Snodgrass and Cross 1983: 15), and the Florida Panhandle (Peck and Thomas 1998: 16).
Three eastern North American species.
There is no published key for the identification of these species.
This species ranges along the Appalachian Mountains from southwestern Virginia (Hoffman et al. 2006: 18) to northern Georgia (Towns County, CMNH).
This taxon has been treated as an aberration of
This species ranges from Nova Scotia (CNC) to northern Indiana (Barr 1969: 74), south to northeastern Georgia (Fattig 1949: 8) along the Appalachian Mountains. The records from northern Florida (Harris 1839: 192) and “Louisiana” (Roeschke 1907a: 145) are probably in error.
This species is known from a few localities in Virginia (Bell 1959: 11), northeastern West Virginia (Hampshire and Randolph Counties, CMNH), the Allegheny Plateau in southwestern Pennsylvania (Allegheny and Fayette Counties, Robert L. Davidson pers. comm. 2008), and the Big Black Mountain in southeastern Kentucky (Davis and Barbour 1978: 139).
Hoffman et al. (2006: 18) reported that distinction between this taxon and
Five eastern North American species.
Van Dyke (1936) reviewed the species. Gidaspow (1973: 78-89) revised the species and provided a key for their identification. One new species was subsequently described by Allen and Carlton in 1988.
This species ranges from Cape Breton Island to southeastern Manitoba, south to eastern Kansas (Popenoe 1877: 22), southeastern Missouri, and Massachusetts [see Gidaspow 1973: Fig. 6]. The record from southeastern Pennsylvania (Rathvon 1869: 524) is likely in error.
This species is known from eastern Kansas, western Missouri, western Arkansas (Allen and Thompson 1977: 32; Allen and Carlton 1988: 136), and central Oklahoma [see Gidaspow 1973: Fig. 6]. The records from “Iowa” and “Texas” (Leng and Beutenmüller 1894: 138) are likely in error; that from Buffalo, New York (Leng and Beutenmüller 1894: 138), is in error.
This species ranges from southeastern Minnesota (Gandhi et al. 2005: 923) south to northern Arkansas (Allen and Thompson 1977: 32; Allen and Carlton 1988: 136), west to eastern Kansas [see Gidaspow 1973: Fig. 6]. The record from “Texas” (Schwarz 1895: 270) is probably in error.
This species is known only from Newton County in northwestern Arkansas.
This species, as far as known, is restricted to Arkansas (Stone County, CNC), western Louisiana, and eastern Texas [see Gidaspow 1973: Fig. 6].
This subgenus includes ten species-group taxa found along the Appalachians. Several undescribed taxa are known (Thomas C. Barr, Jr. pers. comm. 2009).
There is no recent systematic treatment of the species of
This subspecies has been recorded so far only from the Black Mountains in southwestern North Carolina. Roeschke (1907a: 160) reported that this form occurs above 5000 feet of altitude.
This subspecies is known along the Appalachian Mountains from western North Carolina (Leng and Beutenmüller 1894: 138), northeastern Georgia (Fattig 1949: 9), and northwestern South Carolina (Ciegler 2000: 32). It is found from 1800 to 5000 feet (Roeschke 1907a: 160).
This species is known only from the holotype collected in southwestern Virginia.
This species is known from the Cumberland Plateau in southeastern Kentucky and the Appalachian Valley in southwestern Virginia (Barr 1969: 76; Barr 2009: 314).
This species is known from two specimens collected in Cumberland and Anderson Counties, eastern Tennessee (Barr 2009: 315).
This species is known along the Appalachian Mountains in eastern Tennessee and western North Carolina (Barr 1969: 76).
This species is known from southwestern Virginia (Hoffman et al. 2006: 18) and the Black Mountains in western North Carolina (Beutenmüller, 1913: 139).
This species ranges from southeastern Ohio and Pennsylvania (Barr 1969: 76) south to western North Carolina (Thomas C. Barr, Jr. pers. comm. 2009; Leng and Beutenmüller 1894: 138).
This species is endemic, as far as known, to the Black Mountains (Barr 1969: 75) in southwestern North Carolina.
Barr (1969: 75) listed this taxon as a subspecies of
This species is found in the Blue Ridge Mountains (Great Balsams and Highlands Plateau) and the Nantahala Mountains (Barr 1969: 75) in southwestern North Carolina and northeastern Georgia.
Seven species restricted to the Appalachian region.
Valentine (1935) reviewed the species and provided a key for their identification.
This species is endemic to the Black Mountains in western North Carolina (Barr 1969: 74). The record from the mountains of Tennessee (Roeschke 1907a: 147) needs confirmation.
This subspecies is found in the mountains of southwestern Virginia, western North Carolina (Valentine 1936: 227), and northeastern Tennessee (Carter County, CMNH).
This subspecies is known only from Orange and Guilford Counties in northern North Carolina (Valentine 1936: 226). The record from “Tennessee” (Roeschke 1907a: 147) needs confirmation.
This subspecies ranges along the Appalachian Mountains from the Great Smokies in eastern Tennessee and western North Carolina eastwards to the Saluda Mountains (Barr 1970: 4) in northwestern South Carolina.
This subspecies is found along the Appalachians in southern West Virginia (Fayette and Webster Counties, CMNH), eastern Kentucky, southwestern Virginia, and northeastern and southeastern Tennessee (Valentine 1936: 231; Davis and Barbour 1978: 139). The records from Pennsylvania (Roeschke 1907a: 148), southwestern Ohio (Wright and Whitehouse 1941: 70), and southern Indiana (Blatchley 1910: 43) probably refer to other subspecies of the species.
This subspecies ranges from southwestern Pennsylvania and southern Ohio to north-central Kentucky (Valentine 1936: 232) and northwestern Maryland (Glaser 1996: 4).
This subspecies is known in the Appalachians from the Nantahala Mountains, adjacent Blue Ridge, and the Unicoi Mountains in southwestern North Carolina, southeastern Tennessee, and northeastern Georgia (Valentine 1936: 227; Barr 1970: 4).
This subspecies is known only from southeastern Kentucky (Pulaski County, CMNH) and the Walden Ridge in southeastern Tennessee (Valentine 1935: 357).
This species is found along the Appalachians from south-central West Virginia (Fayette County, CMNH) and Virginia (Valentine 1935: 363) to northern Georgia (Rabun, Towns, and Union Counties, CMNH) and northwestern South Carolina (Ciegler 2000: 31).
This subspecies is found from central Tennessee (Barr 1969: 74) to west-central Alabama (Löding 1945: 11).
This subspecies is known only from eastern Alabama (Löding 1945: 11).
This subspecies is known from the Appalachian Plateau in southwestern Pennsylvania and the Allegheny Mountains in West Virginia (Barr 1969: 75), western Virginia (CMNH), and northwestern Maryland (Bailey et al. 1994: 320).
This subspecies is known from northwestern West Virginia and a few counties in northern and western Virginia (Valentine 1935: 367). The record from “Tennessee” (Roeschke 1907a: 150) needs confirmation.
This species is found at high altitudes in the Great Balsam, Plott Balsam, Great Smoky (Barr 1969: 74), and Nantahala Mountains (Barr 1970: 5) in eastern Tennessee, western North Carolina, and northeastern South Carolina.
This species has been reported from the Appalachians in western North Carolina, northern Georgia (Valentine 1935: 358), and northwestern South Carolina (Ciegler 2000: 32).
Four western North American species.
Gidaspow (1973: 73-78) revised the species and provided a key for their identification.
This species is restricted to southeastern Washington (Gidaspow 1973: Fig. 4) and northeastern Oregon (Hatch 1953: 46; Westcott et al. 2006: 9).
This species is known from western Montana (Russell 1968: 42), northern Idaho, Whitman County in southeastern Washington (CMNH), and the Creston area (Lindroth 1961a: 20) in southeastern British Columbia [see Gidaspow 1973: Fig. 4].
This species is known from southeastern British Columbia, Idaho, and eastern Washington [see Gidaspow 1973: Fig. 5].
This form was listed as a synonym of
This species is found in southern British Columbia, western Alberta, western Montana (Russell 1968: 42), northern Idaho, eastern Washington, and from one isolated locality in southwestern Oregon [see Gidaspow 1973: Fig. 5]. Old specimens simply labeled from California are also known (Gidaspow 1973: 77).
Four species restricted to the Pacific coastal and western montane regions of North America.
Van Dyke (1944) and Gidaspow (1973) revised the species then placed in this subgenus. Gidaspow (1973) provided a key for their identification.
This species ranges from Kodiak Island in the Gulf of Alaska to northwestern California, east at least to eastern British Columbia [see Gidaspow 1973: Fig. 2]. The record from Torrington, Alberta (Gidaspow 1973: 68) is probably based on a mislabeled specimen.
This species is known only from Waldo Lake area in western Oregon (Gidaspow 1973: 66; CNC, CMNH).
This rarely collected species is restricted to the southern part of Vancouver Island (Lindroth 1961a: 22) and the Olympic Mountains in northwestern Washington (Van Dyke 1924b: 3).
This species has been placed in the subgenus
This species ranges from northwestern Oregon to the San Francisco Bay area in California [see Gidaspow 1973: Fig. 3]. The record from Vancouver Island (LeConte 1869c: 370) is probably in error.
Fifteen western North American species of which two extend into the Baja California Peninsula.
Gidaspow (1968) revised the species and provided a key for their identification.
This species is found within the Sierra Nevada in central California, from El Dorado County to Sequoia National Park [see Gidaspow 1968: Fig. 9].
This species is restricted to central California [see Gidaspow 1968: Fig. 7].
This species ranges from Sonoma County in California to the Pacific Coast of Baja California Norte, east to the San Bernardino Mountains [see Gidaspow 1968: Fig. 10]. One specimen from Hidalgo state in Mexico is probably mislabeled, as pointed out by Gidaspow (1968: 176).
This species ranges from southwestern Oregon south to the Los Angeles region in southern California, east to the northern part of the Sierra Nevada [see Gidaspow 1968: Fig. 2].
The range of this species extends over much of California, including the Coast Ranges and the Sierra Nevada, as far south as Riverside County [see Gidaspow 1968: Fig. 6]. Some specimens simply labeled from Oregon are known.
The range of this species extends from the Aleutians Islands and the Gulf Coast of Alaska south to northern California, northern Arizona (Coconino County, Eric van den Berghe pers. comm. 2009), and northwestern Wyoming [see Gidaspow 1968: Fig. 4]. Gidaspow (1968: 149) considered the possibility that the species ranges further east toward the Hudson Bay because of the presence in collections of several specimens labeled from “Hudson Bay Territory.”
This species is known from the northern part of California, as far south as Madera County [see Gidaspow 1968: Fig. 5], and from Washoe County in northwestern Nevada (La Rivers 1947: 133, as
This species is known from central California [see Gidaspow 1968: Fig. 3] and from Grand County in eastern Utah (Gidaspow 1968: 165). The record from “Nevada” (Bousquet and Larochelle 1993: 84) needs confirmation.
This species is found in the southern third of California, including Santa Catalina Island, and in “Baja California” [see Gidaspow 1968: Fig. 7].
This species is found in the southern half of California, in Tulare (Sequoia National Park), Fresno, Kern, and Los Angeles Counties [see Gidaspow 1968: Fig. 5].
This subspecies is found in southwestern Oregon (Westcott et al. 2006: 9) and northwestern California as far south as Colusa County [see Gidaspow 1968: Fig. 3].
This subspecies is known for sure only from western Oregon as far south as the border with California [see Gidaspow 1968: Fig. 3]. The records from “California” (Casey 1897: 328, 332, as
This species is found along western California from Mendocino County south to Los Angeles County [see Gidaspow 1968: Fig. 9].
This species is known only from central California, mostly along the west side of the Sierra Nevada from Calaveras County to Tulare and Kern Counties [see Gidaspow 1968: Fig. 8].
This species ranges from southern Oregon to San Luis Obispo County along the coast and to Yosemite National Park along the Sierra Nevada; also known from the Santa Catalina Island [see Gidaspow 1968: Fig. 10].
Three species restricted to the Pacific coastal region of North America.
Van Dyke (1944) first reviewed the species. Subsequently, Gidaspow (1973) revised them and provided a key for their identification.
This species ranges from southwestern British Columbia, including Vancouver Island, to southwestern Oregon [see Gidaspow 1973: Fig. 1].
The range of this species extends near the Pacific Coast from Lincoln County in Oregon (Gidaspow 1973: 61) to Humboldt County in northern California [see Gidaspow 1973: Fig. 1].
This species is known only from a few specimens collected in Humboldt and Mendocino Counties (Weber and Kavanaugh 1992: 394), northern California.
Worldwide, with about 1,080 species arrayed in three genera:
Worldwide, with about 170 species in the Nearctic (41 species, of which one is adventive), Neotropical (about 55 species, many shared with North America), Australian (three species), Oriental (six species), Palaearctic (about 45 species), and Afrotropical (about 35 species) Regions. The species are arrayed in about 25 genus-group taxa.
Gidaspow (1959) revised the North American species and provided a key for their identification. Subsequently Lindroth (1961a: 50, 55) listed in synonymy some of the species that were considered valid by Gidaspow (e.g.,
The genus
Burgess and Collins (1917: 86) reported that
Western Hemisphere, with 14 species in the Nearctic (one species) and Neotropical (14 species) Regions.
This species ranges from Long Island, New York (Notman 1928: 209) to “Iowa” (Burgess and Collins 1917: 62), south to Guatemala (Gidaspow 1963: 301) and southern Florida except for the Keys (Peck and Thomas 1998: 15), west along southern United States to “California” (Burgess and Collins 1917: 62); also known from the Greater Antilles as far south as Puerto Rico (Gidaspow 1963: 301). The records from “Minnesota,” “North Dakota,” and “Wisconsin” (Bousquet and Larochelle 1993: 70) are probably in error or based on strays.
Northern Hemisphere, with six species in the Nearctic (two species, one of them adventive) and Palaearctic (five species) Regions.
This subgenus is retained in its narrow sense and excludes members of
This species occurs from Cape Breton Island to the Skeena River drainage in west-central British Columbia (Lindroth 1961a: 47), south to northeastern Nevada (La Rivers 1947: 134), central Utah (La Rivers 1947: 135), southeastern Texas in the Galveston area (Snow 1906a: 140; Gidaspow 1959: 245), and northern Georgia (Fattig 1949: 11).
This European species was introduced as early as 1906 in many places in Canada and United States for the biological control of two introduced lymantriids: the gypsy moth,
Western Hemisphere, with four species in the Nearctic (four species, one of them endemic) and Neotropical (three species) Regions.
This subgenus is listed in synonymy with the nominotypical subgenus by some authors (e.g., Breuning 1927: 155; Lorenz 2005: 68).
This species is found from the Rio Grande in southeastern Texas (Wickham 1897: 102; Burgess and Collins 1917: 122) south to Nicaragua (Gidaspow 1963: 283) [see Gidaspow 1959: Fig. 2].
This widely distributed species ranges from western Maine (Majka et al. 2011: 45) to northeastern North Dakota (Tinerella 2003: 635), including southern Quebec (only as strays) and Ontario (Lindroth 1961a: 46), south to Venezuela (Gidaspow 1963: 283) and southern Florida (Peck and Thomas 1998: 15), west along the southwest to “California” (Gidaspow 1959: 250) and Baja California (Leng 1915: 565). Two specimens, likely strays, have been collected in New Brunswick (Webster and Bousquet 2008: 16) and on Sable Island, Nova Scotia (Majka et al. 2007: 6). The species is not known from the West Indies.
This species is known from northeastern Georgia and southern Florida but only from a few century-old specimens (Gidaspow 1959: 248), and from the Bahamas (Turnbow and Thomas 2008: 12), Cuba, and Hispaniola (Gidaspow 1963: 283). Erwin (2007a: 104) listed this species also from Yucatán, Mexico.
This species ranges from southeastern New Hampshire (Rockingham County [probably only as strays], Donald S. Chandler pers. comm. 1992) to southeastern Minnesota (Donald P. Schwert pers. comm. 1989), including southern Quebec (only as strays) and southern Ontario (Lindroth 1961a: 47), south to southeastern Texas (Brazoria County, Brian Raber pers. comm. 2010), southeastern Louisiana (Saint John the Baptist and Saint Tammany Parishes, Igor M. Sokolov pers. comm. 2009), and central Georgia (Fattig 1949: 11). The record from “California” (Burgess and Collins 1917: 38) is likely in error or based on a stray.
Three North American species, all of them extending into Mexico.
This subgenus is listed in synonymy with
This species ranges from “Iowa” (Jaques and Redlinger 1946: 295, as
This species is found from central California to eastern Utah, south to southwestern New Mexico, Sonora in Mexico, and Baja California (Gidaspow 1959: 256).
This species is found from Inyo County in eastern California (Riley 1893: 239; Fall 1901a: 40) to central New Mexico, south to Sonora and the Baja California Peninsula (Gidaspow 1959: 255).
Western Hemisphere, with five species in the Nearctic (four species, one of them endemic) and Neotropical (four species, only one,
This species is found in Mexico south at least to Colombia and Venezuela (Erwin 1991a: 26). It is also occasionally found in southwestern United States from California to Texas (Gidaspow 1959: 253).
This species is found in southern California and northern Baja California (Gidaspow 1959: 259); it is also known from one locality in southwestern New Mexico (Gidaspow 1959: 259), from Montezuma County in Colorado (FFPC), and has been reported from Nevada by Erwin (2007a: 91).
The range of this species extends from Sacramento County in central California (Clark 1999: 202) to western Kansas (Popenoe 1877: 22), south to the Federal District in Mexico and Baja California (Gidaspow 1959: 258).
This species is known from western Utah (Millard County, Ken Karns pers. comm. 2009), western Nevada, southern California, and “Arizona” (Gidaspow 1959: 260).
Three North American species, two of them extending into Mexico.
This species ranges from “Vermont” (Ross T. Bell pers. comm. 1989; probably only as strays) to “Nebraska,” including southernmost Ontario (only as strays), south to “Texas” (Gidaspow 1959: 274) and southern Georgia (Fattig 1949: 10).
This species is known from “Arkansas” (Jeannel 1940: 214) to “New Mexico” (Erwin 2007a: 98), south to northeastern Mexico (Gidaspow 1959: 274) and “Louisiana” (Burgess and Collins 1917: 25).
This species is known from “Colorado” (Gidaspow 1959: 275) and southeastern Arizona (Snow 1906b: 161) south at least to the state of Guerrero in Mexico (Gidaspow 1959: 275). The records from southern Kansas (Snow 1903: 192; Knaus 1905a: 218; Burgess and Collins 1917: 26) are doubtful (see Gidaspow 1959: 275).
Seven Mexican species, one of them extending into North America.
This subspecies is known from southern Colorado (Wickham 1902: 231), northwestern Arizona, “New Mexico,” Brewster County in western Texas (Gidaspow 1959: 281), and Chihuahua in Mexico (Erwin 2007a: 96).
This subspecies is known from Mexico as far south as the Federal District and from Brewster County in western Texas (Gidaspow 1959: 281).
Western Hemisphere, with ten species in North America (nine species) and Middle America (four species, only one,
This species ranges from southern Minnesota (Gandhi et al. 2005: 922) to the Pacific Coast in southern California, south to the state of Oaxaca in Mexico (Gidaspow 1959: 269). The records from Missouri (probably based on the type locality of
The range of this species extends from Newfoundland (Larson and Langor 1982: 592) and Saint Pierre and Miquelon (Lindroth 1955a: 28) to eastern British Columbia (Lindroth 1961a: 51), south to “Oregon” (Hatch 1953: 52), northeastern New Mexico (Casey 1897: 344), “Kansas” (Horn 1872c: 384), east-central Missouri (Summers 1873: 133), and northern Georgia (Fattig 1949: 11); several specimens have also been caught along the Slave and Mackenzie Rivers, up to 65°30'N, in Northwest Territories (White 1851: 357; Lindroth 1961a: 51). The records from southeastern Louisiana (Summers 1874a: 79), “Alabama,” “Arkansas,” and “Mississippi” (Bousquet and Larochelle 1993: 72) are probably in error.
Gidaspow (1959: 266) treated
This species ranges from south-central British Columbia (Lindroth 1961a: 53) to north-central North Dakota (McHenry County, Foster F. Purrington pers. comm. 2009), south to “Arizona” (Gidaspow 1959: 263) and southern California along the coast (Casey 1913: 64-65, as
This species inhabits the Great Plains ranging from the southern parts of the Prairie Provinces south to Montana (Gidaspow 1959: 265; Lindroth 1961a: 51) and north-central South Dakota (Walworth County, CNC). The record from “Wyoming” (Bousquet and Larochelle 1993: 73) needs confirmation.
This species is known from southern California to “Colorado,” south to Durango in western Mexico (Gidaspow 1959: 270).
This species ranges from Saskatchewan (Lindroth 1961a: 50) to eastern Oregon, south to northeastern Nevada (La Rivers 1947: 136), northwestern New Mexico (Casey 1897: 345, as
This species ranges from eastern Oregon and “Idaho” (Gidaspow 1959: 271) south to southern Arizona (Snow 1907: 141) and southern California along the coast (Fall 1901a: 40; Moore 1937: 4); also found on Guadalupe Island, Mexico (Gidaspow 1959: 271). The record from northeastern Kansas (Popenoe 1877: 22) is likely in error.
This species is confined to California where it is known from Yolo County to Riverside (Gidaspow 1959: 272) and San Diego Counties (Moore 1937: 4). The records from “Arizona,” “Colorado,” “Texas,” and “Mexico” (Burgess and Collins 1917: 93) are probably in error.
This species is found from Vancouver Island (Lindroth 1961a: 52) to “Nebraska,” south to “Colorado” (Gidaspow 1959: 264) and southern California (La Rivers 1947: 136). The records from “North Dakota” (Bousquet and Larochelle 1993: 73) and “Alberta” (Burgess and Collins 1917: 107) need confirmation.
Obydov (2003: 536), based on a study of two specimens only, concluded that
Fourteen North American species in the western half of the continent.
This species is known only from the original 16 specimens collected at the type locality in the Sierra Nevada.
This species has been recorded so far from the Sierra Nevada in Tulare County and “South Fork” in Humboldt County, California (Gidaspow 1959: 308).
This species is found in California from El Dorado and Sacramento Counties to Santa Cruz and Madera Counties (Gidaspow 1959: 308); it was cited also from Washington (Burgess and Collins 1917: 119) but the record is probably in error or based on a stray.
According to La Rivers (1947: 137), this species is “apparently the predominant
This species is found in California from Sacramento County to Los Angeles and San Bernardino Counties (Gidaspow 1959: 307); according to Burgess and Collins (1917: 122), it was also collected in Reno, western Nevada.
This species ranges from southern Saskatchewan to the Okanagan Valley in south-central British Columbia (Lindroth 1961a: 56), south to southeastern California, northwestern New Mexico (Casey, 1913: 75, as
Gidaspow (1959) considered
This species ranges from Saskatchewan to southeastern British Columbia (Lindroth 1961a: 53-54), south to “California,” “Arizona,” and “Nebraska” (Gidaspow 1959: 305). The records from “Vancouver Island” (Gidaspow 1959: 305) and “Northwest Territory, Canada” (Burgess and Collins 1917: 114) are probably in error.
The range of this species extends from central Washington south at least to El Dorado County in California and western Nevada (Gidaspow 1959: 315); also recorded from “Utah” (Erwin 2007a: 79). One specimen simply labeled from Wyoming is known (Gidaspow 1959: 315). Notwithstanding Gidaspow (1959: 315), Hatch (1953: 54) did not record this species from Washington, Idaho, Oregon, and British Columbia and so the records from “Oregon” and “Idaho” (Bousquet and Larochelle 1993: 74) need confirmation.
This species is known from four specimens without locality data from “Oregon” (Gidaspow 1959: 317).
This species is confined to California where it is found as far south as San Diego County along the coast and Tuolumne County in the Sierra Nevada (Gidaspow 1959: 310).
This species ranges from western Oregon (Hatch 1953: 54, as
This species is known from “Washington,” “Idaho,” and “California” (Burgess and Collins 1917: 114; Gidaspow 1959: 305). The records from British Columbia (Hatch 1953: 53; Gidaspow 1959: 305) probably refer to
This rarely collected species is known from Klamath and Harney Counties in southern Oregon and Siskiyou County in northern California (Gidaspow 1959: 316); also recorded from “Nevada” (Erwin 2007a: 82).
This species is found west of the Rocky Mountains from south-central British Columbia south to “California” (Lindroth 1961a: 54).
About 910 species (Lorenz 2005: 72-114) in North America (14 species), Mexico (two species), and the Palaearctic (about 895 species) and Oriental (about five species) Regions.
Van Dyke (1944) reviewed the North American species and provided a key for the identification of all species except
Twenty-six species (Deuve 2004: 109-119) in the Nearctic (three species, one of them adventive) and Palaearctic (24 species) Regions.
Members of
This European subspecies is adventive in North America where it is known from Newfoundland (Larson and Langor 1982: 592) to southeastern Manitoba (Roughley et al. 2010: 230; CMNH), and from east-central Minnesota (Gandhi et al. 2011: 673), Massachusetts (Van Dyke 1945a: 129), and Connecticut (Middlesex County, William L. Krinsky pers. comm. 2012) in the east, and from western British Columbia (Lindroth 1961a: 37), including the Queen Charlotte Islands (Kavanaugh 2010: 385), western Washington (Hatch 1953: 51), and Edmonton, Alberta (UASM), in the west. The first inventoried specimen collected in the east was found in New Brunswick in 1890 (Lindroth 1961a: 37) and in the west in Seattle, Washington, in 1924 (Hatch 1933c: 117).
The range of this species extends from southern Maine (Dearborn and Donahue 1993: 2) to southeastern Minnesota (Donald P. Schwert pers. comm. 1989), including southernmost Ontario (Lindroth 1961a: 35), south to northeastern Mississippi (Snodgrass and Cross 1983: 14), northern Alabama (Löding 1945: 11), and northern Georgia (Fattig 1949: 10; CMNH). The record from Idaho (LeConte 1878a: 471; Hatch 1953: 51) is in error (Lindroth 1961a: 35); that from southeastern Louisiana (Summers 1874a: 79) needs confirmation.
This species ranges from Massachusetts (Harris 1833: 567; Frost 1910: 86) to southeastern Minnesota (Gandhi et al. 2005: 923), including southern Ontario (Lindroth 1961a: 34), south to southeastern Mississippi (Stone County, UASM), southwestern Alabama (Van Dyke 1945a: 126), and the Florida Panhandle (Peck and Thomas 1998: 16). The record from “Louisiana” (Summers 1874a: 79) is probably in error.
Eleven species in North America (one northern species) and Asia (ten species, one of them extending into northern European Russia).
Deuve (2004: 186) included members of this taxon in the subgenus
This species occurs from the Labrador coast to the Seward Peninsula in Alaska (Lindroth 1961a: 33-34), including the Aleutians and Kodiak Island, south to northern British Columbia (CNC); isolated on high mountains in Gaspé Peninsula (Quebec), New Hampshire (Lindroth 1961a: 34), and Maine (Mount Katahdin, CNC). Fossil remnants of this species, dated between about 16,700 and 21,500 years B.P., have been unearthed in southeastern Iowa (Baker et al. 1986: 96) and northeastern Illinois (Garry et al. 1990: 394).
Thirteen species (Deuve 2004: 197-203) in North America (one Holarctic species) and Asia (13 species, one of them extending into northern European Russia).
This Holarctic subspecies is known from a few old specimens collected on the Kamchatka Peninsula (Shilenkov in Kryzhanovskij et al. 1995: 43) and from Alaska, including the Pribilof Islands, to the Mackenzie River delta in northern Northwest Territories (Lindroth 1961a: 40).
1. This species is listed in the subgenus
One Holarctic species.
This Holarctic subspecies ranges from western Siberia (Bousquet et al. 2003: 154) to the Bering Sea Coast and from Alaska (Lindroth 1961a: 36) to Newfoundland (Lindroth 1955a: 23), south at least to northeastern New York (Powell 1977: 148), northern Iowa (Larsen et al. 2003: 292), and northern New Mexico (Brantley et al. 2003: 382; Sandoval County, UASM). The record from “Pennsylvania” (Bousquet and Larochelle 1993: 76) needs confirmation.
Two other subspecies of this species,
Four species in North America (one species), Asia (two species), and Europe (one species).
This species ranges from Newfoundland (Lindroth 1955a: 26) to the Okanagan Valley in south-central British Columbia (Lindroth 1961a: 41), south to eastern Oregon (Hatch 1953: 50), central New Mexico (Fall and Cockerell 1907: 156), eastern Kansas (Popenoe 1877: 22; Horn 1872c: 384), and northwestern South Carolina (Ciegler 2000: 29).
Ten species in Europe and the Middle East (Deuve 2004: 167-173), one of them adventive in North America.
This European subspecies is adventive in North America where it is found from Newfoundland (Lindroth 1955a: 27) to central Minnesota (Crow Wing County, CNC), south to northeastern Virginia (Falls Church, UASM) in the east, and from the Queen Charlotte Islands (Kavanaugh 1992: 51) to central Alberta (Lindroth 1961a: 37), south to southeastern Wyoming (Burne 1989: 290), northern Utah (Salt Lake County, CMNH), and central California (Lindroth 1961a: 37); seemingly isolated in the Saskatoon area, Saskatchewan (Ronald R. Hooper pers. comm. 1990). The first inventoried specimens collected on this continent was found in New Brunswick in 1890 (Horn 1892d: 61) and on the west coast around 1909 in Seattle, Washington (Hatch 1933c: 117). The date of 1870 listed by Lindroth (1961a: 37) for the first occurrence of this species on the continent is probably a lapsus for 1890. The species was also intentionally introduced in New England as predators of gypsy moths prior to 1911 (Smith 1959: 9).
Four European species, one of them extending into Siberia. One species is adventive in eastern North America.
The European
This European subspecies was intentionally introduced into Massachusetts in 1908 and 1910 (Smith 1959: 7) for gypsy moth control. It is now established in New England (Lindroth 1961a: 37). The first inventoried specimen collected subsequently to its release was found in June 1920 in Winchester, Massachusetts (Smith 1959: 7).
In a cladistic analysis conducted by Arndt et al. (2003: 312, Fig. 7.16), this species turned out as the sister-group to all remaining species of
Five species in North America (four species) and Mexico (two species). One of Mexican species,
This species is known from Kansas (Chautauqua County, CMNH; Erwin 2007a: 110), Wichita National Forest in Oklahoma, and eastern Texas (Van Dyke 1945a: 117).
This species is known from southeastern Arizona and the state of Durango in Mexico (Van Dyke 1945a: 118-119).
This species ranges from “Maine” (Larochelle and Larivière 1990a: 27) to northern Minnesota (Gandhi et al. 2005: 923), north to north-central Ontario (Spires 1985: 79), south to “Texas” (Van Dyke 1945a: 116, as
This subspecies ranges from Newfoundland (Lindroth 1955a: 25) to Yukon Territory, south to the Sierra Nevada in east-central California, southern Arizona, southern New Mexico, and southwestern South Dakota (Purrington et al. 2002: 202) [see Edelbrock 1986: Fig. 69].
This subspecies is known from Vancouver Island and the Puget Sound area in Washington south to central Oregon between the coast and the Cascade Range (Edelbrock 1986: 49, Fig. 69).
This subspecies is known from high altitude along the northern Cascades and from coastal mountains of Oregon (Edelbrock 1986: 49).
This subspecies is restricted to southeastern Alaska including the Aleutian Islands (Edelbrock 1986: 49, Fig. 69).
1. Edelbrock (1986) studied the geographical variation in
Twenty-two species (Deuve 2004: 363-375) in North America (one Holarctic species) and Eurasia (22 species).
This subspecies is found in eastern Siberia and in North America from the Seward Peninsula in Alaska to Bathurst Inlet on the arctic coast of Nunavut (Lindroth 1961a: 42).
Two other subspecies of this species occur in eastern Asia.
Worldwide, with more than 2,500 species (Lorenz 2005: 22-62) arrayed in five tribes:
Western Hemisphere, with 13 species in North America (ten species), Mexico (two species), and South America (one species) arrayed in three genera:
Seven species (Pearson et al. 2006: 48) in western North America (five species) and northern Mexico (two species:
Vaurie (1955) reviewed the species and offered a key for their identification. Subsequently one new North American species,
The record of
This species, also known as the “Montane Giant Tiger Beetle
This species, also known as the “Great Plains Giant Tiger Beetle”, ranges from southwestern South Dakota (Howden 1970: 8) and eastern Wyoming (Huber 1978: 75) south to western Texas [see Pearson et al. 2006: Map 3]. The record from “Arizona” (Horn 1910a: 123) is in error; those from “Arkansas” (Leng 1902: 97; Erwin and Pearson 2008: 20) are probably based on a misinterpretation of the type locality.
This species, also known as the “South Texas Giant Tiger Beetle”, is found only in south and west-central Texas (Pearson et al. 2006: 51).
This species, also known as the “Plateau Giant Tiger Beetle”, occurs from the northern half of Arizona to western Texas, north to southwestern Colorado (Kippenhan 1994: 21) and southeastern Utah (Krell and Brookhart 2012: 110). The record from “Oklahoma” (Bousquet and Larochelle 1993: 52) is in error.
This species, also known as the “Mojave Giant Tiger Beetle”, is found from northwestern Arizona and southwestern Utah west to the desert mountains of southeastern California east of the Sierra Nevada [see Vaurie 1955: Fig. 1 and Pearson et al. 2006: Map 2].
Five species in western North America inhabiting the Pacific coastal lowlands and the mountain slopes, including those of the Cascade and Sierra Nevada ranges.
Casey, between 1897 and 1924, described so many forms in this genus (90 species-group taxa) that at the time of his death in 1925 it was virtually impossible to identify members of
The range of this species, also known as the “Audouin’s Night-stalking Tiger Beetle”, extends from southwestern British Columbia, including southern Vancouver Island, south to northwestern California [see Pearson et al. 2006: Map 9].
This subspecies, also known as the “Narrow Night-stalking Tiger Beetle”, is found above 1500 m of elevation in Plumas and Lassen Counties in northeastern California (Leffler 1979a: 218).
This subspecies, also known as the “California Night-stalking Tiger Beetle”, ranges from southwestern Oregon to southern California along the coast and through the Sierra Nevada, at elevation below 900 m near the range of
This subspecies, also known as the “Intermediate Night-stalking Tiger Beetle”, is found above 900 meters in the Sierra Nevada between El Dorado and Tulare Counties, California (Leffler 1979a: 222).
This subspecies, also known as the “Subcylindrical Night-stalking Tiger Beetle”, is restricted to San Martin, Santa Clara County, in western California (Leffler 1979a: 217).
This taxon, also known as the “Mount Ashland Night-stalking Tiger Beetle”, is known only from Jackson County in southwestern Oregon.
This species, also known as the “Greater Night-stalking Tiger Beetle”, is found from southwestern British Columbia, including Vancouver Island, to southwestern Oregon [see Pearson et al. 2006: Map 6]. The record from northern California (Leng 1902: 105) needs confirmation; that from “Montana” (Leng 1902: 105) is in error.
This species, also known as the “Lustrous Night-stalking Tiger Beetle”, is restricted to a small area east of Warthan Canyon in western Fresno County, central California (Knisley and Haines 2010: 245).
About 200 species in the Nearctic (four species), Neotropical (about 165 species), Australian (25 species), Palaearctic (one species), and Afrotropical (12 species) Regions. The species are arrayed in 11 genera (see Naviaux 2007: 10-11):
Western Hemisphere, with about 95 species (110 species-group taxa) (Naviaux 2007: 3) in the Nearctic (four species, only one endemic) and Neotropical (about 95 species) Regions. The species are arrayed in seven subgenera:
Naviaux (2007) recently revised the species and provided keys for the subgenera and the species groups but not for all species. Pearson et al. (2006: 24) field guide to the tiger beetles of North America included a key to all North American species (
In their phylogenetic analysis based on the nuclear 18S and the mitochondrial 16S and cytochrome oxidase III genes, Zerm et al. (2007) found out that the genus
From a zoogeographical point of view it is of interest to note that a specimen identical to present day
Twenty-nine species in North America (three species, one of them endemic) and the Neotropical Region (28 species), including the West Indies.
The range of this subspecies, also known as the “Pan-American Big-headed Tiger Beetle” (Pearson et al. 2006: 57) or the “Carolina Metallic Tiger Beetle” (Erwin and Pearson 2008: 70), extends from Maryland to southeastern California, south to Nicaragua (Naviaux 2010: 70) and the Florida Keys [see Pearson et al. 1997: Fig. 3]; two specimens from San Diego County in California are also known (Moore 1937: 3). The records from the Bahamas, Cuba, Cayman Islands, and Jamaica refer to
Three more subspecies are recognized by Naviaux (2007: 40):
This species, also known as the “Florida Metallic Tiger Beetle”, is found only in southern Florida, from Dixie County to the Keys (Choate 2003: 63).
The range of this species, also known as the “Virginia Big-headed Tiger Beetle” (Pearson et al. 2006: 59) or the “Virginia Metallic Tiger Beetle” (Erwin and Pearson 2008: 100), extends from southwestern Connecticut (Dunn 1985b: 21) to southwestern Nebraska (Spomer et al. 2008a: 54), south to southwestern Texas, northeastern Mexico (Pearson et al. 2006: 60), and the Florida Keys [see Pearson et al. 2006: Map 13]; apparently isolated in southeastern Arizona (Dajoz 2004: 116) and south-central Wisconsin (Lawton 1971: 57). The records from “Colorado” (Boyd 1982: 5) and “Tabasco” (Erwin and Pearson 2008: 100) need confirmation.
About 55 Neotropical species, one of them extending into southeastern Texas.
This species, also known as the “Upland Metallic Tiger Beetle”, ranges from southeastern Texas (Darlington 1935b: 161, as
This species has passed under the name
Worldwide, with more than 1,710 species arrayed in five subtribes:
Worldwide, with about 1,090 species. The North American fauna is represented by 98 species (about 9% of the world fauna). The number of genera recognized varies greatly depending on the approach used (lumper versus splitter).
Willis (1968) published a simplified key based on external characters for 91 North American forms, representing all the current recognized species except for the following 11:
About 210 species in all zoogeographical regions. The North American fauna includes eight species (about 4% of the world fauna) placed in the nominotypical subgenus.
Lorenz (2005: 54-58) listed the following taxa as subgenera of
About 45 species (Lorenz 2005: 55-56) in the Nearctic (eight species, 14 species-group taxa), Neotropical (about 15 species), Oriental (four species), Palaearctic (14 species), and Afrotropical (six species) Regions.
This species, also known as the “Swift Tiger Beetle”, has been recorded from Nebraska and western Iowa south to north-central Texas and north-central Arkansas [see MacRae and Brown 2011b: Fig. 8]. The record from Indiana (Montgomery and Montgomery 1931: 359) was based on misidentified
The range of this species, also known as the “Ant-like Tiger Beetle”, extends from North Dakota and west-central Minnesota (Tinerella and Rider 2000: 367) to the Ohio River Valley in West Virginia (Kirchner and Kondratieff 1999: 84), south to western Alabama (Löding 1945: 10), Louisiana, and northern Kansas (Popenoe 1877: 22) [see Pearson et al. 2006: Map 90]; also recorded from “Montana” (Erwin and Pearson 2008: 207).
This species, also known as the “Grass-runner Tiger Beetle”, ranges from southeastern Arizona to southwestern Texas [see Pearson et al. 2006: Map 87], south to Durango (Cazier 1954: 287).
This subspecies, also known as the “White-striped Tiger Beetle”, ranges from southern California to southwestern New Mexico, as far north as southwestern Utah (Tanner 1929a: 86), south to Nayarit (Cazier 1960: 12) [see Shook 1989: Fig. 1].
This subspecies, the “Rouged Tiger Beetle”, is known from northeastern New Mexico south to southern Durango and Tamaulipas [see Shook 1989: Fig. 1].
Intergrades between the two subspecies of
This species, also known as the “Meadow Tiger Beetle”, is restricted to the western and northern slopes of the Sierra Nevada and adjacent eastern portion of the San Joaquin Valley of California (Woodcock et al. 2006: 869). Specimens labeled from Riverside in California, Prescott in Arizona (see Cazier 1939: 28), and Olmito and San Benito in Texas are doubtful or in error (Woodcock et al. 2006: 869); the records from San Pedro Mártir Mountains in Baja California (Cazier 1948: 18) and “Nevada” (Boyd 1982: 16) need confirmation. Extensive surveys in the past 20 years have produce a sole extant population, near Westwood in Lassen County leading Woodcock et al. (2006: 875) to suggest that the species should be considered as a candidate for listing as a Threatened and Endangered species by the U.S. Fish and Wildlife Service.
This form has been considered a subspecies of
The range of the “Belted-winged Tiger Beetle” is disputed. According to Johnson (1990b: Fig. 1), it ranges from west-central Alberta to southeastern Saskatchewan, south to central New Mexico and central Arizona; isolated in southwestern Utah and southwestern Yukon Territory. According to Pearson et al. (2006: 153), it is found in “lower elevations of Colorado, New Mexico, and Arizona.” Erwin and Pearson (2008: 222) recorded the subspecies from the same states as Pearson et al. (2006: 153) and added Kansas (also cited by Leng 1902: 155).
This subspecies, the “Interior Tiger Beetle”, is known from Nye County in western Nevada and from the Traverse Mountain Range in western Ventura County eastwards to the San Bernardino Mountains in San Bernardino County and the southern tip of the Sierra Nevada in Kern County, California (Kippenhan 2007: 14, Fig. 8). The record from “NE” (Erwin and Pearson 2008: 222) is probably an error for “NV.”
Freitag (1999: 87) listed this taxon as a junior synonym of
This subspecies, the “Imperfect Tiger Beetle”, ranges from westernmost Alberta and British Columbia, as far north as Fort Saint John along the Peace River (Catling 2007: 19), south to southern Utah, southern Nevada, and Mono County (Kippenhan 2007: 13) in west-central California [see Johnson 1990b: Fig. 1]; also recorded from “Arizona” and “Wyoming” by Erwin and Pearson (2008: 222). The record from New Mexico (Fall and Cockerell 1907: 155) must be in error.
This subspecies, the “Kaibab Tiger Beetle”, is known only from the Kaibab Plateau of northern Arizona (Johnson 1990b: 1).
This subspecies, the “Susan’s Tiger Beetle”, is restricted to the valleys east of the Sierra Nevada Mountains in Mono and Inyo Counties, California [see Kippenhan 2007: Fig. 8].
This subspecies, also known as the “Variable Tiger Beetle”, ranges from northwestern Ontario (Lawton 2008: 73) to southeastern Saskatchewan, south to northwestern Nebraska [see Johnson 1990b: Fig. 1]. The record from Minnesota (Horn 1928: 12) needs confirmation.
The range of this species, also known as the “One-spotted Tiger Beetle”, extends from northeastern New York to central Georgia and southern Mississippi, west to western Missouri [see Pearson et al. 1997: Fig. 19]. The record from “Texas” (Horn 1915: 389) is probably in error or based on a stray.
This subspecies, the “Pygmy Tiger Beetle”, ranges from central Arizona (Pearson et al. 2006: 156) south to Durango (Cazier 1954: 285).
The subspecies
Thirteen North American species (27 species-group taxa), of which five extend into Mexico, as far south as the state of Yucatan, and one into the Bahamas and Cuba.
This species, also known as the “Sandbar Tiger Beetle”, is endemic to the Coastal Plain ranging from southeastern North Carolina to the Florida Panhandle west to southeastern Mississippi and southwestern Louisiana [see Pearson et al. 2006: Map 97].
This species, also known as the “Coppery Tiger Beetle”, ranges from southern Ohio to southwestern Manitoba and central Montana, south to east-central New Mexico, eastern Texas, and southern Alabama [see Pearson et al. 2006: Map 100]; also known from one locality in northern Georgia. The record from “Wisconsin” (Bousquet and Larochelle 1993: 68) is likely in error (Peter W. Messer pers. comm. 2007).
This species, also known as the “Whitish Tiger Beetle”, inhabits the Coastal Plain ranging from Virginia (Hoffman et al. 2006: 17) to the Florida Panhandle, west to southwestern Alabama (Löding 1945: 10) [see Pearson et al. 2006: Map 106]; also recorded from “Mississippi” (Erwin and Pearson 2008: 229). The record from Louisiana (Boyd 1982: 17) is probably in error or based on a stray.
This subspecies, the “Gulf Beach Tiger Beetle”, is found along the Gulf coast of Florida, including the Keys (Choate 2003: Map 61), and Alabama (Löding 1945: 10). The record from Horn Island, Mississippi (Richmond 1968: 234), is in error for
The nominotypical subspecies is known from the states of Tabasco and Veracruz and
This subspecies, the “Coastal Tiger Beetle”, is found along the Gulf Coast from Mississippi (Graves and Pearson 1973: 187) to the state of Veracruz in Mexico (Erwin and Pearson 2008: 230).
This species, also known as the “Moustached Tiger Beetle”, ranges from east-central Georgia (Beaton 2008: 42) to southern Florida (Choate 2003: Map 63; Pearson et al. 2006: 174). The record from “North Carolina” (Erwin and Pearson 2008: 230) is probably in error or based on a stray.
This species, also known as the “Ghost Tiger Beetle”, ranges from southern Quebec to southeastern Alberta (Hilchie 1985: 333), south to Chihuahua (Cazier 1954: 297), central Texas, southern Louisiana, and eastern North Carolina, west to western Arizona and eastern Nevada [see Pearson et al. 1997: Fig. 17]. The record from South Carolina, based on a specimen in CMNH, is probably in error (Knisley and Schultz 1997: 113). According to Erwin and Pearson (2008: 231), the Ghost Tiger Beetle has been extirpated over much of its former range due to habitat loss.
This subspecies, the “Denton Tiger Beetle”, is known from Dallas, Denton, and Kaufman Counties in northern Texas (Vaurie 1951: 10) [see Willis 1967: Fig. 137].
This subspecies, the “Panhandle Tiger Beetle”, is found in Oklahoma, northern Texas, and northeastern New Mexico [see Pearson et al. 2006: Map 101]. The records from “Arkansas” and “Kansas” (Boyd 1982: 17) need confirmation.
Willis (1967) indicated the presence of narrow zones of intergradation between this subspecies and the other two.
This subspecies, also known as the “Sandy Stream Tiger Beetle”, ranges from Beaver Islands in northern Michigan (Dunn 1987: 11) to southern Wyoming, south to north-central Colorado (Kippenhan 1990: 314), Arkansas, and Tennessee [see Pearson et al. 2006: Map 101]. The record from “Ohio” (Willis 1967: 269) needs confirmation (see Graves and Brzoska 1991: 28); that from “Texas” (Freitag 1999: 94) probably refers to the
Intergrades between this and the
This species, also known as the “Margined Tiger Beetle”, is found along the Atlantic Coast from Kings County in Nova Scotia (Neil and Majka 2008: 4) to the Florida Keys and along the Gulf Coast in Florida [see Pearson et al. 2006: Map 96]; it is also recorded from the Bahamas and Cuba (Peck 2005: 27). According to Dunn (1983: 4), the species has declined significantly along the New Hampshire coast.
This species, also known as the “Puritan Tiger Beetle”, is now restricted to two areas, one along the Connecticut River in southern Massachusetts and Connecticut, the other one along the Chesapeake Bay in Maryland (Pearson et al. 2006: 170, 193). It formerly occurred along the Connecticut River as far north as central New Hampshire.
This species has been listed as threatened by the U.S. Fish and Wildlife Service in August 1990.
This species, also known as the “White-sand Tiger Beetle”, is found along a small area within the Coastal Plain from south-central Georgia to eastern Louisiana [see Pearson et al. 2006: Map 98]. Beaton (2008: 42) did not find this species at any historical sites in Georgia during his survey of the tiger beetles of the state.
This subspecies, the “Chiracahua Tiger Beetle”, is known from southeastern Arizona and Sonora, Mexico (Spomer 2004: 409); also recorded from “New Mexico” (Erwin and Pearson 2008: 233).
This subspecies, the “Knaus’s Tiger Beetle”, ranges from southern Manitoba to southeastern Alberta, south to central New Mexico and central Texas [see Willis 1967: Fig. 143].
This subspecies, also known as the “Salt Creek Nevada Tiger Beetle”, has been reported yet only from around the type locality in eastern Nebraska [see Willis 1967: Fig. 143]. Population estimates vary yearly from a few hundred to under a thousand specimens (Spomer et al. 2008a: 43).
This species has been listed as endangered by the U.S. Fish and Wildlife Service in October 2005.
This subspecies, the “Indian Creek Tiger Beetle”, is known only from the type locality in the South Dakota Badlands (Spomer 2004: 410).
This subspecies, also known as the “Nevada Tiger Beetle”, is found in the Great Basin region of Nevada and California south to northern Sonora [see Willis 1967: Fig. 143].
Another subspecies,
This subspecies, the “Olmos Creek Tiger Beetle”, ranges from southeastern Arizona to southeastern Texas, including northern Coahuila [see Willis 1967: Fig. 143].
This subspecies, the “Tube Tiger Beetle”, ranges from northeastern Utah to central Arizona and northwestern New Mexico [see Willis 1967: Fig. 143].
This species, also known as the “Aridland Tiger Beetle”, is found from northern Utah to southern Colorado, south to Chihuahua (Cazier 1954: 297) [see Pearson et al. 1997: Fig. 29]. The record from “Arkansas” (Boyd 1982: 17) is in error or based on a stray.
Rumpp (1977: 176) recognized
This subspecies, the “Inquisitor Tiger Beetle”, is found in central Texas (Pearson et al. 2006: 171).
This subspecies, also known as the “Rio Grande Tiger Beetle”, ranges from northern Utah to northeastern New Mexico [see Pearson et al. 2006: Map 103], south to Durango and Tamaulipas (Cazier 1954: 296). The records from western Kansas (Popenoe 1877: 22; Snow 1878: 63) and “Oklahoma” (Boyd 1982: 17) need confirmation; that from Yuma, California (Leng 1902: 167) is likely in error.
Another subspecies,
Three species (five species-group taxa) in United States and Cuba (one species) and Mexico (two species), including Baja California.
This species, also known as the “Olive Tiger Beetle”, is found in Cuba and southern Florida, including the Keys [see Woodruff and Graves 1963: Fig. 3]. Apparently the species has not been sighted in Florida since the 1980s (Pearson et al. 2006: 149).
Six Neotropical species, of which one is represented by a distinct subspecies in southeastern Texas.
This subspecies, the “Smyth’s Beach Tiger Beetle”, is known only from the type locality in southeastern Texas. No specimens have been collected since the original ones (over 80 specimens) in June 1912.
The nominotypical subspecies is found along the Gulf Coast of Mexico, as far south as Veracruz, and from Honduras (Erwin and Pearson 2008: 290).
Thirty-eight Neotropical species (Lorenz 2005: 58), of which one extends into southwestern North America.
The Cuban
This species, also known as the “Sonoran Tiger Beetle”, ranges from southern Arizona south to Sinaloa; it is also found in southern Baja California Peninsula (Cazier 1954: 286). The record from “California” (Erwin and Pearson 2008: 118) needs confirmation.
Four North American species, one of them extending into northern Mexico.
This species, also known as the “Loamy-ground Dromo Tiger Beetle”, ranges from central and coastal Texas (Pearson et al. 2006: 160) south to Tamaulipas (Cazier 1954: 297); it was also recorded from southeastern Colorado (Michels et al. 2008).
This species, also known as the “Cajun Dromo Tiger Beetle”, is found along and near the Gulf Coast in southern Louisiana (Schaupp 1884a: 85) and northeastern Texas [see Pearson et al. 2006: Map 92].
This species, also known as the “Frosted Dromo Tiger Beetle”, ranges from northern Kansas and central Missouri (MacRae and Brown 2011a) south to Nueces and Webb Counties in southern Texas and west into the Texas Panhandle (Pearson et al. 2006: 160). There is also one record from east-central Nebraska (Pearson et al. 2006: 160).
This species, also known as the “Velvet Dromo Tiger Beetle”, is found only in southern Texas, primarily along the coast [see Pearson et al. 2006: Map 94].
Western Hemisphere, with 18 species (33 species-group taxa) in the Nearctic (nine species, 18 species-group taxa) and Neotropical (14 species) Regions.
This subspecies, the “California Tiger Beetle”, ranges from the Mojave Desert of southern California (Pearson et al. 2006: 138) south to northeastern Baja California Peninsula and northwestern Sonora (Cazier 1954: 289).
This form intergrades with the
This subspecies, the “Inland Tiger Beetle”, is found only in the Death Valley in Inyo and San Bernardino Counties, California (Rumpp 1958: 151).
The nominotypical subspecies is found in the Baja California Peninsula and
This subspecies, also known as the “Cream-edged Tiger Beetle”, is found from southeastern Oklahoma to northeastern Mexico [see Johnson 1993b: Fig. 1].
This form intergrades with the
This subspecies, the “Johnson’s Tiger Beetle”, ranges from central Missouri to southeastern Colorado, north to west-central Nebraska (Spomer et al. 2008a: 58), south to New Mexico and southwestern Texas [see Johnson 1993b: Fig. 1]. The taxon is also found in Coahuila in northern Mexico (Murray 1979: 55). The records from “North Dakota” (Boyd 1982: 14; Freitag 1999: 77) probably refer to the
This subspecies, the “
This subspecies, also known as the “Eastern Beach Tiger Beetle”, once occurred along the Atlantic Coast from Cape Cod, Massachusetts to the Chesapeake Bay. It is now found at two isolated sites on the coast of Massachusetts and along both shores of the Chesapeake Bay in Maryland and Virginia (Boyd and Rust 1982: 234; Pearson et al. 2006: 144). The subspecies was successfully reintroduced at Sandy Hook, New Jersey (Pearson et al. 2006: 192). The records from “Delaware” (Bousquet and Larochelle 1993: 65) and from near Lancaster, Pennsylvania (Cresson 1861: 12) are probably based on strays.
This subspecies is listed as threatened under the Endangered Species Act by the U.S. Fish and Wildlife in 1990 (Pearson et al. 2006: 191).
This subspecies, the “Eastern Beach Tiger Beetle”, is found along the Atlantic Coast from Ocean County in New Jersey to southern Florida (Boyd and Rust 1982: 234).
According to Knisley and Schultz (1997: 103), the ranges of this form and of the
This subspecies, the “Saulcy’s Beach Tiger Beetle”, ranges from the Gulf Coast of Florida to the Mississippi River delta in Louisiana (Pearson et al. 2006: 144); it is also known from the Isla Arcas in Campeche (Bates 1884: 260). The records from Cuba (Leng and Mutchler 1916: 697, as
This subspecies, the “Gulf Beach Tiger Beetle”, is found from coastal Mississippi (Lago et al. 2002: 201) south along the Gulf to the state of Tamaulipas (Cazier 1954: 294). The record from east-central Colorado (Snow 1877: 16) is probably in error.
This subspecies, the “Pale Tiger Beetle”, is known from the Salt Basin of western Texas and adjacent New Mexico and from a single site in Dawson County, Texas (Acciavatti 1981: 239).
This subspecies, the “Willcox Tiger Beetle”, is known only from the type locality in southeastern Arizona (Acciavatti 1981: 238).
This subspecies, also known as the “Glittering Tiger Beetle”, is found from east-central Arizona to north-central New Mexico, south to the Rio Grande area in westernmost Texas and southeastern New Mexico [see Acciavatti 1981: Fig. 1]; also recorded from Chihuahua in Mexico (Murray 1979: 55).
This subspecies intergrades with the
This species, also known as the “Western Tidal Flat Tiger Beetle”, occurs along the Pacific Coast from southern California to central Baja California Peninsula, and along the Gulf of California Coast from northern Sonora to Sinaloa (Cazier 1954: 291) [see Pearson et al. 1997: Fig. 28]. According to Pearson et al. (2006: 139), this species is now found in the United States only in three or four protected areas in Ventura, Orange, and San Diego Counties.
This species, also known as the “Gulfshore Tiger Beetle”, is found along the Gulf Coast from eastern Mississippi (Grammer 2009) to northern Tamaulipas in Mexico (Pearson et al. 2006: 145).
This subspecies, the “Virgin River Tiger Beetle”, is found only along the Virgin River in southwestern Utah and southeastern Nevada [see Acciavatti 1981: Fig. 1].
This subspecies, also known as the “Riparian Tiger Beetle”, is found from the Gila River Basin in eastern Arizona westwards to the Salton Sea Basin in California, north to southern Nevada [see Pearson et al. 2006: Map 74]. The record from New Mexico (Fall and Cockerell 1907: 155) needs confirmation. According to Erwin and Pearson (2008: 251), the species has been extirpated from many of its historic sites.
This subspecies, also known as the “Saltmarsh Tiger Beetle”, is found along the Gulf Coast from the Florida Keys to Tamaulipas in Mexico (Cazier 1954: 261) [see Pearson et al. 1997: Fig. 4]; also recorded from Yucatán (Horn 1897a: 354).
This species, also known as the “Elusive Tiger Beetle”, is found along the Atlantic Coast from southern South Carolina (Cartwright 1935: 73; Ciegler 1997: 191) to central Florida, and along the Gulf Coast of Florida [see Pearson et al. 1997: Fig. 10].
One North American species which extends into northern Mexico.
This subspecies, the “Salt Flat Tiger Beetle”, is known only from Torrance and Guadalupe Counties in central New Mexico and Hudspeth County in western Texas (Pearson et al. 2006: 148).
This subspecies, the “Alkali Tiger Beetle”, ranges from eastern Nebraska (Carter 1989: 15) and central Colorado (Kippenhan 1990: 312) south to northern Texas (Gaumer and Murray 1971: 10) and southeastern New Mexico (Acciavatti et al. 1980: 31) [see Pearson et al. 2006: Map 80]. The record from north-central Utah (Tanner 1929a: 87) is probably in error.
This subspecies, also known as the “White-cloaked Tiger Beetle”, is known from scattered localities from southern South Carolina (Cartwright 1935: 75) to northeastern Texas, south to northern Florida (Choate 2003: Map 48) and Tamaulipas (Cazier 1954: 297) [see Pearson et al. 2006: Map 80]. Ciegler (1997: 191) reported that the last specimen seen from South Carolina was collected in 1935 and that the species may be extinct in the state.
Worldwide, with about 340 species described by 2005 assigned to 24 subgenera (Lorenz 2005: 41-51). The North American fauna is represented by 60 species (about 18% of the world fauna) placed in two subgenera.
The genus is employed here in a restricted sense as used by most taxonomists working on the Palaearctic and African faunas and recently by Erwin and Pearson (2008) for the North American fauna.
Western Hemisphere, with about 65 species (Lorenz 2005: 48-49) in the Nearctic (21 species, 42 species-group taxa) and Neotropical (about 55 species).
1.
This species, also known as the “Eastern Pinebarrens Tiger Beetle”, is found mainly along the Coastal Plain and Piedmont Plateau from Long Island in southeastern New York (Leng 1928: 206) to central Florida, west to southeastern Louisiana [see Pearson et al. 1997: Fig. 11].
This subspecies, also known as the “Great Basin Tiger Beetle”, is found in the Death Valley area in eastern California (Leffler 1987: 8).
1. Rumpp (1956: 141) reported the presence of intergrade populations between this subspecies and the
This subspecies, also known as the “Nye Tiger Beetle”, is found in southeastern Oregon (Leffler 1979a: Fig. 60) and western Nevada (Rumpp 1956: 140).
Kippenhan (2005) indicated from an analysis of populations that the variation in the dorsal coloration in
This species, also known as the “Cazier’s Tiger Beetle”, is found along a small area in Jim Hogg and Starr Counties, southeastern Texas [see Pearson et al. 2006: Map 63], and in Tamaulipas, Mexico (Erwin and Pearson 2008: 127).
This taxon is listed as a subspecies of
This species is known only from a few sites in the Richmond Heights area of Miami (Brzoska et al. 2011: 5).
This subspecies, also known as the “Grand Canyon Tiger Beetle”, is restricted to the Colorado River at the bottom of the Grand Canyon in northern Arizona and along the Virgin River in adjacent Utah and Nevada (Pearson et al. 2006: 135).
The range of this subspecies, also known as the “Wetsalts Tiger Beetle”, extends from central Washington to northwestern Wyoming, south to western Texas and along the Pacific Coast to the northern parts of the Baja California Peninsula (Cazier 1948: 11) [see Pearson et al. 1997: Fig. 35].
1. Some authors (e.g., Nagano 1982: 39) have treated
This species, also known as the “Highlands Tiger Beetle”, is found in Highlands and Polk Counties, central Florida (Choate 2003: 84; Pearson et al. 2006: 126).
This species, also known as the “Horn’s Tiger Beetle”, ranges from southern Arizona to southwestern Texas, south to Durango (Cazier 1954: 248) [see Pearson et al. 1997: Fig. 25].
The subspecies
The range of this species, also known as the “Cobblestone Tiger Beetle”, is disjunct: one population is known from New Brunswick (Sabine 2005: 53) south to central New Jersey (Boyd 1978: Fig. 28), northwestern West Virginia (Allen and Acciavatti 2002: 26), southeastern Kentucky (Laudermilk et al. 2010: 28), and southeastern Indiana; the second is found in northeastern Mississippi and western Alabama [see Pearson et al. 2006: Map 67]. The record from “South Carolina” (Choate 2003: Map 30) needs confirmation.
This species is listed on the IUCN Red List of Threatened Species (IUCN 2007) and has been extirpated from many historical sites (Erwin and Pearson 2008: 155).
This subspecies, the “Bowditch’s Tiger Beetle”, is known from southwestern Colorado (Kippenhan 1994: 65) and northwestern New Mexico (Rumpp 1962: 172). The record from “Arizona” (Boyd 1982: 11) is in error or based on a stray.
This subspecies intergrades with the nominate form in north-central New Mexico (Pearson et al. 2006: 119).
This subspecies, also known as the “Black Sky Tiger Beetle”, occurs from Salton Sea in southern California (LaRue 1991: 49) and southernmost Nevada (Kippenhan 2002: 381) to western Texas (Gaumer and Murray 1971: 10), north to southwestern Utah (Tanner 1929a: 85) and northeastern Colorado (Kippenhan 1990: 311), south to Aguascalientes and San Luis Potosí in Mexico (Cazier 1960: 8).
This subspecies, the “Subtropic Tiger Beetle”, is confined to Hidalgo and Cameron Counties in southern Texas (Pearson et al. 2006: 119).
This subspecies, the “Rio Grande Grassland Tiger Beetle”, is known from the lower Rio Grande Valley in southern Texas from Maverick County to Hidalgo County (Pearson et al. 2006: 122), north to Kimble County in central Texas (Mawdsley 2009: 9).
This subspecies, also known as the “Large Grassland Tiger Beetle”, ranges from western Kansas to central Arizona, north to northern Colorado (Kippenhan 1994: 66), south to southern New Mexico and southwestern Texas; also known from one locality in eastern Kansas [see Mawdsley 2009: Fig. 18]. The record from “Utah” (Boyd 1982: 11) is likely in error.
This subspecies intergrades with the
This subspecies, the “Santa Clara Grassland Tiger Beetle”, ranges from southern Colorado (Kippenhan 1994: 67) south to northern Durango (Cazier 1954: 251), including southwestern Texas and western Arizona [see Mawdsley 2009: Fig. 18].
This subspecies, the “Prairie Tiger Beetle”, ranges from southern Missouri and north-central Arkansas to north-central Texas, south to southeastern Texas [see Mawdsley 2009: Fig. 18] and Coahuila in Mexico (Cazier 1954: 250). The record from central New Mexico (Fall and Cockerell 1907: 154) is suspect (Mawdsley 2009: 8); that from “Colorado” (Wickham 1902: 228) is probably in error.
This subspecies, also known as the “Ocellated Tiger Beetle”, occurs from southeastern Arizona and adjacent New Mexico (Pearson et al. 2006: 133) south to Costa Rica (Blackwelder 1944: 18); also recorded from “Texas” (Erwin and Pearson 2008: 162).
This subspecies, the “Dark-abdomened Tiger Beetle”, ranges from western Louisiana (Graves and Pearson 1973: 180) to northern New Mexico [see Pearson et al. 2006: Map 66], including southern Oklahoma (Schmidt 2004: 5), south to southern Tamaulipas (Cazier 1954: 278).
This subspecies, the “Barbaraann’s Tiger Beetle”, occurs in the Hueco, Sierra Diablo, and Apache mountains in western Texas (Gage 1988: 146-147) and in the Sacramento Mountains of southern New Mexico (Pearson et al. 2006: 129) where it is found above 1500 m.
This subspecies, the “Rock-loving Tiger Beetle”, is known only from above 1670 m in the Guadalupe Mountains in western Texas and southeastern New Mexico (Gage 1988: 146).
This subspecies, also known as the “Limestone Tiger Beetle”, occurs from Carter and Murray Counties in southern Oklahoma (Pearson et al. 2006: 128) south to Coahuila and Nuevo León (Sumlin 1985: Fig. 9).
Another subspecies,
This subspecies, the “Green Mountain Tiger Beetle”, is known only from the type locality in southeastern New Mexico.
This subspecies, the “Chihuahua Tiger Beetle”, is known from northeastern Colorado to west-central Nevada [see Pearson et al. 2006: Map 56], south to Chihuahua (Cazier 1954: 253). The records from Oklahoma (Drew and Van Cleave 1962: 113), “Nebraska,” “Kansas,” and “Texas” (Freitag 1999: 68) apparently refer to intergrades and these records are registered under the nominotypical subspecies.
Bertholf (1983: 21) listed this form in synonymy with the nominotypical subspecies.
This subspecies, also known as the “Punctured Tiger Beetle”, ranges from New Brunswick to southern Alberta, south to southern Texas and southern Florida [see Pearson et al. 2006: Map 56].
The two subspecies of
This subspecies, also known as the “Tascoa Tiger Beetle”, has been recorded from Oldham County in northwestern Texas (Davis 1918: 34), Alajuela province in Costa Rica (Horn 1905: 22), and Panama (Erwin and Pearson 2008: 178).
1. No specimens of this subspecies have been collected in United States since the original ones in 1917 and the subspecies has never been found in Mexico. Cazier (1954: 279) believed that the US specimens of
This subspecies, the “Mexican Red-bellied Tiger Beetle”, ranges from Louisiana (Pearson et al. 2006: 131) south through Texas to Queretaro and Veracruz (Murray 1979: 53). The records from “Georgia” (Boyd 1982: 12) and western Alabama (Löding 1945: 9) probably refer to the nominotypical subspecies; those from “Mississippi,” “Arkansas” (Boyd 1982: 12), and Oklahoma (Drew and Van Cleave 1962: 114) are probably based on intergrades and these records are listed under the nominotypical subspecies.
This subspecies, the “Hentz’s Tiger Beetle”, is found only along eastern Massachusetts [see Leonard and Bell 1999: Fig. 111]. The record from “Rhode Island” (Bousquet and Larochelle 1993: 64) is in error or based on a stray.
This subspecies, also known as the “Eastern Red-bellied Tiger Beetle”, ranges from southwestern Vermont (Leonard and Bell 1999: 104) to southwestern Missouri, south to east-central Texas and the Florida Panhandle [see Pearson et al. 2006: Map 64].
This subspecies intergrades with the
This species, also known as the “Scabrous Tiger Beetle”, is found from southeastern Georgia to southern Florida [see Pearson et al. 2006: Map 61].
This species, also known as “Schaupp’s Tiger Beetle”, occurs from southeastern Kansas and northwestern Arkansas [see Pearson et al. 2006: Map 69] south to Nuevo León (Cazier 1954: 287). The record from “Missouri” (Erwin and Pearson 2008: 181) needs confirmation.
This subspecies, also known as the “Western Red-bellied Tiger Beetle”, ranges from northeastern New Mexico to central Arizona [see Pearson et al. 2006: Map 65], south to Guanajuato in Mexico (Cazier 1954: 271).
Besides the nominotypical subspecies, four other subspecies are known, ranging collectively from Mexico to Costa Rica (see Pearson et al. 2006: 132; Erwin and Pearson 2008: 185-186).
This species, also known as the “Senile Tiger Beetle”, is found along western California, as far north as Sonoma and Lake Counties, and the northern part of the Baja California Peninsula. According to Pearson et al. (2006: 116), it is now known in the United States only from a few protected coastal populations and two interior populations, one near Lake Elsinore in western Riverside County and one near Jacumba in San Diego County.
This species, also known as the “Thin-lined Tiger Beetle”, ranges from western Nebraska (Brust 2007: 9) to southern California [see Pearson et al. 1997: Fig. 38], south to northern Sinaloa and southern Tamaulipas (Cazier 1954: 257).
This highly vagile subspecies, also known as the “Ascendant Tiger Beetle”, is found primarily along the Atlantic and Gulf Coasts from Virginia to southern Florida, west to eastern Texas [see Pearson et al. 2006: Map 70], south to Panama (Erwin and Pearson 2008: 197). This form has also been found inland as far north as north-central Kansas (Charlton and Kopper 2000: 266) and along the Atlantic Coast as far north as Massachusetts (Comboni and Schultz 1989: 151); however there is no known established populations inland anywhere in North America (Pearson and Vogler 2001: 105). The record from the “West Indies” (LeConte 1856a: 51) needs confirmation.
Besides the two subspecies found in North America, six other subspecies are recognized among this polymorphic species in Middle and South America and the West Indies.
This subspecies, the “Sigmoid Tiger Beetle”, is found along the Pacific Coast from southern California (Nagano 1982: 38) to the Baja California Peninsula (Cazier 1954: 291); it is occasionally found inland, as far as the Salton Sea in Imperial County (see LaRue 1991). The record from “Arizona” (Freitag 1999: 74) needs confirmation.
This subspecies, the “Echo Tiger Beetle”, is found mainly within the Great Basin from Wyoming to southern Oregon, south to east-central California, southern Nevada, and southern Utah (Pearson et al. 2006: 114).
This subspecies, the “Torrance Tiger Beetle”, is known only from Torrance County in central New Mexico (Pearson et al. 2006: 114).
This subspecies, the “Funaro’s Tiger Beetle”, is known only from the type locality in northwestern New Mexico.
This subspecies, the “Hairy-fronted Tiger Beetle”, is found in central Kansas, western Oklahoma, west-central Texas, and east-central New Mexico (Willis 1967: 302); also recorded from “Arizona” (Erwin and Pearson 2008: 202).
This subspecies, the “Augured Tiger Beetle”, is known from Inyo County in eastern California and Nye County in Nevada (Rumpp 1956: 135).
This subspecies, the “Owens Lake Tiger Beetle”, is restricted to Owens and adjacent Panamint Valley of east-central California (Pearson et al. 2006: 115).
This subspecies, the “Sulphur Valley Tiger Beetle”, is endemic to the Sulphur Springs Valley in southeastern Arizona.
This subspecies, also known as the “Williston’s Tiger Beetle”, is endemic to the Laramie Plain of Wyoming (Rumpp 1962: 168).
This subspecies intergrades with the
Northern Hemisphere, with about 75 species (Lorenz 2005: 43-48) in the Nearctic (38 species, of which four extend into northern Mexico; 92 species-group taxa) and Palaearctic (36 species) Regions.
This eastern species, also known as the “Appalachian Tiger Beetle”, ranges from southwestern New Brunswick (Webster and Bousquet 2008: 16) south to northeastern Georgia, west at least to eastern Kentucky (Laudermilk et al. 2010: 28) [see Pearson et al. 1997: Fig. 15]. Old records from Indiana, northern Illinois, and western Missouri (see Wilson and Larochelle 1980: 37-38) suggest that the species was more widely distributed at one time. Beaton (2008: 40) did not find the species in Georgia during his intensive survey of tiger beetles in the state.
This species, the “Death Valley Tiger Beetle”, is known from the Death Valley region (Kritsky and Horner 1998: 17) in Inyo County, California, and Nye County, Nevada.
This form has been listed as a subspecies of
This subspecies is restricted to the area of ancient Lake Bonneville in north-central Utah [see Knisley et al. 2012: Fig. 23].
This subspecies, also known as the “Badlands Tiger Beetle”, ranges in patchy colonies from eastern Alaska south through the Rocky Mountains to northeastern New Mexico and southern Utah, east to western North Dakota and western Nebraska [see Knisley et al. 2012: Figs 23, 24]. The records from “Kansas” (Leng 1902: 134) and “Manitoba” (Knisley et al. 2012: 14) need confirmation.
This subspecies ranges from south-central British Columbia to southeastern Washington [see Knisley et al. 2012: Fig. 23].
This subspecies is restricted to high elevations of the Bear River Mountains of southeastern Idaho and northeastern Utah [see Knisley et al. 2012: Fig. 23].
This species, also known as the “Green Claybank Tiger Beetle”, inhabits the Great Plains from eastern Montana and North Dakota south to northern Louisiana, northern Texas, and northeastern New Mexico [see Schincariol and Freitag 1991: Fig. 13].
According to Pearson et al. (2006: 92), individuals with green elytra and blue head and thorax from northwestern Louisiana and southwestern Arkansas (originally described under the name
This subspecies, also known as the “Crimson Saltflat Tiger Beetle”, ranges from Minnesota to southern Alberta (Hilchie 1985: 330), south to northeastern Arizona and northern Texas [see Pearson et al. 1997: Map 11]. The record from “Saskatchewan” (Freitag 1999: 27) needs confirmation.
Pearson et al. (2006: 99), followed by Erwin and Pearson (2008: 138), considered
This subspecies, also known as the “Alkaline Tiger Beetle”, is found in southern Wyoming and northwestern Colorado (Kippenhan 1996a: 42).
Pearson et al. (2006: 99) recorded this subspecies from a much larger area, throughout the western Great Plains and intermontane southern Rocky Mountains.
This subspecies, also known as the “Westbourne’s Tiger Beetle”, is found in southern Manitoba, southern Saskatchewan (Wallis 1961: 51), north-central North Dakota, and northwestern Minnesota (Knudsen 1985: 186); also recorded from “Montana” (Erwin and Pearson 2008: 139).
This subspecies, also known as the “Western Beach Tiger Beetle”, is found along the Pacific Coast from southern California to the central parts of the Baja California Peninsula (Cazier 1948: 14) and also along the Gulf of California coast in northern Sonora [see Pearson et al. 1997: Fig. 27]. According to Pearson et al. (2006: 195), this taxon is now gone from most of its former sites in southern California.
Some authors (e.g., Nagano 1982: 37) consider
This subspecies, the “Jorda’s Tiger Beetle”, is known from northeastern Arizona (Bertholf 1983: 12) and northern New Mexico (Acciavatti et al. 1980: 30). Based on Pearson et al. (2006: Map 47), it is also found in southern Utah and southwestern Colorado. The record from “Wyoming” (Boyd 1982: 9) probably refers to the nominotypical subspecies.
This subspecies, also known as the “Blowout Tiger Beetle”, ranges from western South Dakota (Spomer et al. 2008a: 57) and southern Wyoming, south to northern New Mexico, extreme northwestern Texas, and southern Oklahoma [see Pearson et al. 2006: Map 47]. The records from “Montana” (Horn 1915: 372) and “Iowa” (Boyd 1982: 9) need confirmation.
This subspecies intergrades with the
This subspecies, the “Adroit Tiger Beetle”, ranges from southern Manitoba to northern Alberta (Wallis 1961: 56), south to Wyoming and northwestern South Dakota (Spomer et al. 2008a: 34).
This species, also known as the “Common Claybank Tiger Beetle”, ranges from Newfoundland to eastern British Columbia, north to northern Yukon Territory (Eagle River, Sydney G. Cannings pers. comm. 2009), south to eastern Utah, northern New Mexico, central Missouri, southern Pennsylvania, and New Jersey [see Schincariol and Freitag 1991: Fig. 13; Pearson et al. 1997: Fig. 3]. According to Knisley and Schultz (1997: 114), the literature records from Virginia, western North Carolina, and northwestern Georgia could refer instead to
Spomer et al. (2008a: 25) noted that this species intergrades occasionally with
This species, also known as the “Autumn Tiger Beetle”, is confined to the Coastal Plain and Piedmont Plateau ranging from “North Carolina” (Knisley and Schultz 1997: 116) to the Florida Panhandle (Choate 2003: Map 13), west to southeastern Mississippi (George County, Drew A. Hildebrandt pers. comm. 2009).
This species, the “Ohlone Tiger Beetle”, is known only from remnant stands of native grassland on coastal terraces in Santa Cruz County, California (Freitag et al. 1993: 117). According to Pearson et al. (2006: 193), it is known from only nine sites where populations range from less than 100 to several hundreds.
This species has been listed as endangered under the Endangered Species Act by the U.S. Fish and Wildlife Service in October 2001. Collection of specimens is illegal (Pearson et al. 2006: 88).
This subspecies, also known as the “Dark Saltflat Tiger Beetle”, occurs from southeastern Oregon and southwestern Idaho to southwestern Utah (Leffler 1987: 7). The record from “Washington” (Freitag 1999: 39) probably refers to the
Some authors, including Freitag (1999: 39), consider
This subspecies, the “Platt Tiger Beetle”, occurs in southeastern Oregon, east-central California, and western Nevada [see Pearson et al. 2006: Map 42].
This subspecies, the “Wallis’ Tiger Beetle”, ranges from south-central British Columbia south to southeastern Oregon (Leffler 1987: 7), including southwestern Idaho (Shook 1984: 159).
This species, also known as the “Cochise Tiger Beetle”, is restricted to southeastern Arizona, southwestern New Mexico [see Pearson et al. 1997: Fig. 31] and adjacent regions in Sonora, Mexico (LeConte 1856a: 62, as
This rare species, also known as the “Alpine Tiger Beetle”, ranges from southern Idaho to south-central Oregon, south to east-central California and west-central Utah [see Pearson et al. 2006: Map 31]. The record from one locality in northern Montana (see Pearson et al. 2006: Map 31) is possibly based on a stray.
This species, also known as the “Sagebrush Tiger Beetle”, is found from southern British Columbia south to north-central Oregon [see Pearson et al. 1997: Fig. 32].
According to Pearson et al. (2006: 89), further studies may show that this taxon is rather a subspecies of
This subspecies, the “Audubon’s Tiger Beetle”, ranges from southern Manitoba to central British Columbia, south to east-central California, Arizona, and northern Texas [see Pearson et al. 2006: Map 28]. The records from “Wisconsin,” “Illinois,” “Kentucky,” “Tennessee” and “Arkansas” (Boyd 1982: 7) apparently refer to the nominate form.
This subspecies, the “Cimarron Tiger Beetle”, is known from north and central Colorado (Kippenhan 1994: 44-45), much of New Mexico (Acciavatti et al. 1980: 30), and southeastern Arizona (Bertholf 1983: 22) [see Pearson et al. 2006: Map 28]. The records from northern (Tanner 1929a: 79) and southwestern (Horn 1926: 266) Utah need confirmation.
This subspecies, also known as the “Hatch’s Tiger Beetle”, ranges from Vancouver Island to the central region of the Sierra Nevada in California (Leffler 1987: 4).
This subspecies intergrades with the
This subspecies, the “Elegant Tiger Beetle”, ranges from the Columbia River in southern Washington to northwestern California (Leffler 1987: 4).
This subspecies, also known as the “Cow Path Tiger Beetle”, ranges from southern Quebec to Minnesota, south to central Arkansas and northern Georgia [see Pearson et al. 2006: Map 28]. Beaton (2008: 39) indicated that despite extensive search he was unable to find any population of this species in Georgia, including at all known historical sites, and Ciegler (1997: 189) noted that it has not been collected in South Carolina since 1936. The record from “Nova Scotia” (Bousquet and Larochelle 1993: 59) was based on a misidentified specimen (see Majka et al. 2007: 6); that from “New Brunswick” (Erwin and Pearson 2008: 175) needs confirmation.
This subspecies intergrades with the
This subspecies, the “Chartreuse Tiger Beetle”, is known only from north-central Texas (Pearson et al. 2006: 102).
This subspecies, the “LeConte’s Tiger Beetle”, ranges from southern Quebec to southern Manitoba (Wallis 1961: 35-36), south to Kansas, northern Mississippi, West Virginia, and Connecticut [see Pearson et al. 2006: Map 43). The records from “Saskatchewan” and “Alberta” (Bousquet and Larochelle 1993: 60) refer to the nominotypical subspecies.
This subspecies intergrades with the nominate form over a broad area in the Great Plains and with the
This subspecies, the “Rugate Tiger Beetle”, is known from southwestern Arkansas (Ward 1972: 70), western Louisiana (Graves and Pearson 1973: 175), and eastern Texas (Vaurie 1950: 4). The record from “Oklahoma” (Erwin and Pearson 2008: 183) needs confirmation.
This subspecies intergrades with the nominate form in the northwestern part of its range and with the
This subspecies, also known as the “Wrinkle-fronted Tiger Beetle”, ranges east of the Appalachians from Massachusetts (Leonard and Bell 1999: 133) to North Carolina (Harris 1911: 28, as
This form intergrades with the
This subspecies, also known as the “Festive Tiger Beetle”, ranges from southwestern Saskatchewan and southeastern Alberta (Wallis 1961: 35) south to east-central New Mexico (Acciavatti et al. 1980: 30), northern Texas (Gaumer and Murray 1971: 10), Arkansas (Graves and Pearson 1973: 175), and northwestern Mississippi (Bolivar County, Drew A. Hildebrandt pers. comm. 2009). The record from “Iowa” (Freitag 1999: 49) probably refers to the
According to Pearson et al. (2006: 102), this subspecies intergrades with the
This subspecies, the “Unicolored Tiger Beetle”, is known from eastern Tennessee and North Carolina (Pearson et al. 2006: 103) south to central Florida (Peck and Thomas 1998: 16; Choate 2003: Map 74), southern Alabama (Löding 1945: 9), and southern Mississippi (Graves and Pearson 1973: 174); also recorded from “Virginia” (Erwin and Pearson 2008: 184). The records from Texas (Tucker 1906: 85), “Louisiana,” “Colorado” (Leng 1902: 125), New Mexico (Fall and Cockerell 1907: 155), and Oklahoma (Drew and Van Cleave 1962: 115) could be based on strays, on intergrades, or be in error.
According to Pearson et al. (2006: 103), this form intergrades with the
This subspecies, the “Yampa Tiger Beetle”, is known only from the Maybell Sand Hills area in northwestern Colorado (Kippenhan 1994: 55).
This species, also known as the “Splendid Tiger Beetle”, occurs from southern Pennsylvania to eastern Wyoming, north to southern Wisconsin, south to central Texas, northeastern Georgia, and northern South Carolina [see Schincariol and Freitag 1991: Fig. 13; Pearson et al. 1997: Fig. 20]; The records from “New York” (Schaupp 1884a: 90), “Minnesota” (Horn 1928: 10), and “New Mexico” (Freitag 1999: 50, 51) need confirmation.
1. The name
This species, also known as the “Short-legged Tiger Beetle”, is found from southern Oregon to east-central California, east to southeastern Utah [see Pearson et al. 2006: Map 48]. The records from “Arizona” and “New Mexico” (Bousquet and Larochelle 1993: 61) are in error or based on strays.
This subspecies is known only from east-central Arizona [see Kritsky and Horner 1998: Fig. 5].
Bertholf (1983: 26) considered this form as a synonym of the
This subspecies, the “Diffracted Tiger Beetle”, is found from southern Wyoming to New Mexico and eastern Arizona (Kritsky and Horner 1998: 25, Fig. 5). According to Pearson et al. (2006: 107), it occurs in southern Nevada, northern Arizona, and New Mexico; also recorded from Utah (Tanner 1929a: 80) and “Nebraska” (Erwin and Pearson 2008: 192).
This subspecies, the “Joaquin Tiger Beetle”, is known only from the San Joaquin Valley of California.
According to Knisley and Haines (2007: 109), this subspecies intergrades with the
This subspecies, the “Kirby’s Tiger Beetle”, ranges from Manitoba to Alberta, north to southern Northwest Territories; it southern limit is debated. Kritsky and Horner (1998: Fig. 5) placed it at southern Wyoming and northern Nebraska while Pearson et al. (2006: Map 45) placed it at southeastern New Mexico and northern Texas. I have accepted the latest range for the records. The subspecies is also recorded from “British Columbia,” “Ontario,” “Arkansas,” “Iowa,” “Minnesota,” “Missouri,” “Utah,” and “Washington” by Erwin and Pearson (2008: 193).
This subspecies intergrades with the nominotypical form over a large area in the Midwest (Pearson et al. 2006: 106).
This subspecies, also known as the “Opal Tiger Beetle”, occurs in Utah, Nevada, and eastern California (Kritsky and Horner 1998: 25).
Some authors, including Pearson et al. (2006: 107, 108), consider the forms
This subspecies, the “Sierra Tiger Beetle”, is restricted to the Sierra Nevada in eastern California [see Kritsky and Horner 1998: Fig. 5].
This subspecies, also known as the “Oblique-lined Tiger Beetle”, ranges over much of eastern North America, from Newfoundland to Nebraska, south to northern Texas and northern Florida (Choate 2003: Map 26) [see Kritsky and Horner 1998: Fig. 5; Pearson et al. 2006: Map 45]. Several state and province records (e.g., AB, CA, ID, MB, MT, ND, NT, NV, OR, SK, UT, WY) in Boyd (1982: 9), Bousquet and Larochelle (1993: 61), and Freitag (1999: 57) apparently refer to other subspecies of
This subspecies, the “Wealed Tiger Beetle”, ranges from British Columbia to western Montana, south to northern Utah and northern California [see Kritsky and Horner 1998: Fig. 5]. The record from “Northwest Territories” (Boyd 1982: 10) is probably in error.
Freitag (1999: 55) and Pearson et al. (2006: 107) considered the form
This taxon, also known as the “Santa Ana Tiger Beetle”, is found in southern California, primarily in Orange and western San Bernardino Counties (Pearson et al. 2006: 108).
This species, also known as the “Laurentian Tiger Beetle”, is restricted to a small area in southeastern Manitoba, northwestern (Lawton 2008: 73) and southwestern Ontario, and adjacent parts of Minnesota (Kaulbars and Freitag 1993a: 307; Pearson et al. 2006: 95); isolated at Manitoulin Island, Ontario (Bouchard et al. 2006: 21).
This subspecies, also known as the “Big Sand Tiger Beetle”, ranges from southern Saskatchewan and southern Alberta south to New Mexico and southern Texas (Gaumer 1977: 188-189); also recorded from “Manitoba” and “Minnesota” (Erwin and Pearson 2008: 135).
Gaumer (1977: 194-195) reported the presence of intergrade populations between this subspecies and the
This subspecies, also known as the “Eastern Sand Tiger Beetle”, ranges from Kings County in Nova Scotia (Neil and Majka 2008: 4) to southern Manitoba, south to northeastern South Dakota (Spomer et al. 2008a: 21), central Louisiana, southern Mississippi, and southern Virginia (Gaumer 1977: 202-203). The records from “Saskatchewan,” “Montana,” “Nebraska” (Erwin and Pearson 2008: 136), “Colorado” (Leng 1902: 136), and “Alabama” (Freitag 1999: 25) need confirmation.
This subspecies, also known as the “Gibson’s Sand Tiger Beetle”, is known from southwestern Saskatchewan (Wallis 1961: 38; Gaumer 1977: 216) and northwestern Colorado (Kippenhan 1994: 41). The record from “Alberta” (Bousquet and Larochelle 1993: 55) is in error; that from “North Dakota” (Freitag 1999: 26) needs confirmation; that from “Utah” (Erwin and Pearson 2008: 136) is probably based on intergrades found along the Green River (see Pearson et al. 2006: 84).
In his unpublished thesis, Gaumer (1977: 219) treated the Colorado population of this subspecies as a distinct subspecies of
This subspecies, also known as the “Reddish-green Sand Tiger Beetle”, is found in southwestern Arkansas, northern Louisiana, and eastern Texas (Gaumer 1977: 210).
This subspecies, also known as the “Mescalero Sand Tiger Beetle”, is found in Terry and Yoakum Counties in northwestern Texas and in Chaves, Eddy, and Roosevelt Counties in eastern New Mexico (Rumpp 1986: 143).
Rumpp (1986: 144) reported the presence of intergrade populations between this subspecies and the nominotypical form in Bailey and Lamb Counties, northwestern Texas, and Quay County, eastern New Mexico.
This subspecies, the “Laurent’s Long-lipped Tiger Beetle”, ranges from north-central Montana to western South Dakota, south to central New Mexico, central Arizona, and southern Nevada [see Spanton 1988: Fig. 39].
The range of this subspecies, also known as the “Boreal Long-lipped Tiger Beetle”, extends from Newfoundland to eastern Alaska, south to southern Alberta, central Minnesota, central Wisconsin, and southern New York [see Spanton 1988: Fig. 39]. Intergrade populations between the three subspecies of
The range of this subspecies, the “Green Long-lipped Tiger Beetle”, extends along the Cascade Range and Sierra Nevada from southwestern British Columbia to east-central California (Spanton 1988: 129, Fig. 39). The records from Colorado (Wickham 1902: 228), “Idaho,” and “Montana” (Boyd 1982: 6) probably refer to the
This species, also known as the “Prairie Long-lipped Tiger Beetle”, ranges from the Fraser River in British Columbia to northwestern Ontario (Lawton 2008: 72), south to west-central Nebraska (Spomer et al. 2008a: 54), northern Colorado, southern Utah, and east-central California [see Spanton 1988: Fig. 40]. The records from “New Mexico” (Freitag 1999: 35) and Minnesota (Horn 1928: 11) need confirmation.
This subspecies, also known as the “Consenta’s Tiger Beetle”, was once found in Long Island and New Jersey but is now restricted to the Pine Barrens region of New Jersey (Mawdsley 2007: 17). Single specimens, possibly strays, are known from Delaware, “Maryland,” and “Pennsylvania” (Mawdsley 2007: 17). The taxon is also recorded from “North Carolina” and “Virginia” by Erwin and Pearson (2008: 166).
This subspecies, also known as the “Northern Barrens Tiger Beetle”, ranges from New Hampshire (Leonard and Bell 1999: 47) to Minnesota, south to northern Alabama (Löding 1945: 9) northeastern Georgia, and northwestern South Carolina [see Kaulbars and Freitag 1993a: Fig. 40]. The species has been collected also at two sites in the Outaouais region in western Quebec and eastern Ontario (Leonard and Bell 1999: 47). The record from New Jersey (Smith 1910: 197) is questionable (see Boyd 1978: 215). Beaton (2008: 39) indicated that he was unable to find any extant populations of this species in Georgia despite intensive searching at historical sites and other areas of suitable habitat.
This subspecies, the “Dorothy’s Tiger Beetle”, in known from southeastern Arizona (Bertholf 1983: 21), New Mexico (Acciavatti et al. 1980: 30), and a small area in western Texas [see Pearson et al. 2006: Map 39].
This subspecies, also known as the “Beautiful Tiger Beetle”, ranges from western South Dakota (Larsen and Willis 2008: 480; Brust 2010: 1) and northeastern Wyoming (Brust 2011: 78) south to northern Texas, New Mexico, and Arizona (Bertholf 1983: 20) [see Pearson et al. 2006: Map 39]; also recorded from Nuevo León (Erwin and Pearson 2008: 172).
This species, also known as the “Six-spotted Tiger Beetle”, ranges from Nova Scotia to the Black Hills in southwestern South Dakota, south to east-central Texas and northern Florida [see Kaulbars and Freitag 1993a: Fig. 39; Pearson et al. 2006: Map 36]. The record from “North Dakota” (Freitag 1999: 50) needs confirmation.
This species, also known as the “Coral Pink Sand Dune Tiger Beetle”, is confined to a small area of less than 400 hectares in the Coral Pink Sand Dunes formation in southwestern Utah [see Johnson 1991: Fig. 22] and is considered an endangered species.
This taxon was listed as a subspecies of
This species, also known as the “St. Anthony Dune Tiger Beetle”, is found in the Snake River Valley of southeastern Idaho [see Shook and Clark 1988: Fig. 1; Pearson et al. 1997: Fig. 22] and in southwestern Montana (Winton et al. 2010: 43).
This subspecies, also known as the “Pacific Coast Tiger Beetle”, ranges along the seacoast from Grays Harbor County in northern Washington to Del Norte County in northern California [see Leffler 1979a: Fig. 30; Pearson et al. 2006: Map 26].
This subspecies, also known as the “Frechin’s Tiger Beetle”, is known only from a small area around Neah Bay in the extreme northwestern Olympic Peninsula, Washington (Pearson et al. 2006: 82).
This species, also known as the “Columbia River Tiger Beetle”, was once found along the Columbia, Salmon, and Snake Rivers in north-central Oregon, south-central Washington, and western Idaho [see Leffler 1979a: Fig. 29; Pearson et al. 1997: Fig. 27]. According to Pearson et al. (2006: 81), it is now known only from a few sites along the Salmon River in Idaho.
This species is listed on the IUCN Red List of Threatened Species (IUCN 2007).
This montane subspecies, also known as the “Dispirited Tiger Beetle”, ranges from southern Alaska south to the Sierra Nevada in east-central California and western Nevada; also found in the Rocky Mountains in southeastern British Columbia, northern Idaho, and western Montana [see Freitag 1965: Fig. 33; Pearson et al. 2006: Map 19].
Intergrade populations are known between the two subspecies of
This subspecies, also known as the “Eureka Tiger Beetle”, inhabits a narrow area along or near the Pacific Coast from northern Washington to northern California [see Freitag 1965: Fig. 33].
This species, also known as the “Twelve-spotted Tiger Beetle”, is found from Northwest Territories and the eastern front of the Rocky Mountains in Alberta to Newfoundland (Lindroth 1955a: 16), south to Alabama, central Texas, and Colorado (Kippenhan 1990: 309) [see Freitag 1965: Fig.17]. The record from Vancouver, British Columbia (Wallis 1961: 22) is possibly based on a mislabeled specimen.
This subspecies, also known as the “Sacramento Valley Hairy-necked Tiger Beetle”, is endemic to a small area within the Sacramento Valley of California (Graves et al. 1988: 660). An extensive survey in 2001-2004 within the known distributional range of the subspecies yield no specimens and Knisley and Fenster (2006) concluded that the subspecies has been extirpated in the late 1980s to early 1990s possibly from the construction of the Oroville Dam on the Feather River in the 1960s (see also Fenster and Knisley 2006).
This subspecies, the “Athabascan Tiger Beetle”, is known only from the Lake Athabasca Sand Dunes of northern Alberta and Saskatchewan (Graves et al. 1988: 667). This is the northernmost and most isolated known population of
This subspecies, also known as the “Colorado River Tiger Beetle”, is restricted to the valley of the Little Colorado River in Navajo County, northeastern Arizona (Graves et al. 1988: 669).
This subspecies, the “Southwest Hairy-necked Tiger Beetle”, ranges from western Colorado and western New Mexico westwards to southeastern California and the northern parts of the Baja California Peninsula [see Graves et al. 1988: Fig. 6]. The record from “Wyoming” (Erwin and Pearson 2008: 143) needs confirmation. According to Pearson et al. (2006: 73), the subspecies is now probably extirpated from many former sites along the Gila River in central and western Arizona.
This subspecies, the “Coulee Tiger Beetle”, inhabits mainly the Columbia-Snake river system (Graves et al. 1988: 671) ranging from southern British Columbia to southern Oregon and Idaho. Its western limit is in Cowlitz County, within 100 km of the coast (Pearson et al. 2006: 73). The record from “Montana” (Erwin and Pearson 2008: 143) needs confirmation.
This subspecies, The “Pacific Hairy-necked Tiger Beetle”, is restricted to the Pacific Coast of northern Baja California and southern California (Graves et al. 1988: 659), as far north as Santa Cruz County (Pearson et al. 2006: 73). According to Erwin and Pearson (2008: 143), this taxon is threatened and has been eliminated over most of its former range.
This subspecies, also known as the “Hairy-necked Tiger Beetle”, ranges east of the Mississippi River from New Jersey to southern Florida (Peck and Thomas 1998: 16), west to the Mississippi River Valley (Graves et al. 1988: 658, Fig. 6) then north to the southern region of the Great Lakes. According to Allen and Acciavatti (2002: 12), this subspecies is considered rare and even endangered throughout the Ohio River drainage and other regions. The records from Oklahoma (Drew and Van Cleave 1962: 110), “Kansas,” and “Nebraska” (Erwin and Pearson 2008: 144) are based on intergrades and are recorded here under the
This subspecies intergrades with the
This subspecies, the “Rhode’s Tiger Beetle”, ranges from Newfoundland to the Lake Superior region in Ontario, south to southeastern Wisconsin (Messer 2010: 33), Michigan, Connecticut, and Rhode Island [see Graves et al. 1988: Fig. 6]. Numerous state and province records (e.g., MB, IA, IL, IN, MN, ND, NE, SD) listed by Erwin and Pearson (2008: 145) are reported here under other subspecies of
This subspecies, the “Shelford’s Tiger Beetle”, inhabits the Great Plains from southern Manitoba to southern Alberta, south to New Mexico and eastern Texas (Graves et al. 1988: 665, Fig. 6). Numerous state records (e.g., AR, AZ, LA, MN, UT) listed by Erwin and Pearson (2008: 145) are reported here under other subspecies of
This subspecies intergrades with the nominate form over a large area west of the Mississippi river, from southern Manitoba to Louisiana (Graves et al. 1988: Fig. 6), and with the
This subspecies, the “Northwest Hairy-necked Tiger Beetle”, is found along the Pacific Coast from northern Washington to northern California (Graves et al. 1988: 672). According to Pearson et al. (2006: 74), it is now extirpated from most historic sites.
This subspecies, the “Hyperboreal Tiger Beetle”, ranges from southern Northwest Territories to east-central Alberta, central Saskatchewan, west-central Manitoba [see Johnson 1991: Fig. 22] and northeastern Manitoba (Woodcock et al. 2011: 118).
This subspecies intergrades with the
This subspecies, the “Labrador Tiger Beetle”, is known only from southern Labrador (Johnson 1991: Fig. 22; Brzoska and Stamatov 2008: 50-51).
This subspecies, also known as the “Sandy Tiger Beetle”, is restricted to a small area enclosing southeastern Wyoming, southern South Dakota, Nebraska, and northeastern Colorado [see Johnson 1991: Fig. 22]. The record from “Kansas” (Boyd 1982: 7) needs confirmation.
This subspecies, the “Nogahabar Tiger Beetle”, is known only from the type locality.
This subspecies, also known as the “Nymphal Tiger Beetle”, ranges from central Alberta to northwestern Minnesota, south to northern South Dakota (Spomer et al. 2008a: 19) and northern Montana [see Johnson 1991: Fig. 22; Pearson et al. 2006: Map 21].
This subspecies, the “Dappled Tiger Beetle”, ranges from eastern Alaska to western Northwest Territories, north to above the Arctic Circle (Brzoska 2008: 65), south to the Queen Charlotte Islands and, along the Rocky Mountains, to central New Mexico [see Freitag 1965: Fig. 18]; also recorded from “Arizona” (Erwin and Pearson 2008: 163) and “Idaho” (Boyd 1982: 7).
This subspecies intergrades with the nominate form in southern British Columbia south, along the crest of the Rocky Mountains, to central Utah, with the
This subspecies, the “Maricopa Tiger Beetle”, ranges from southern California to south-central New Mexico, north to southern Nevada [see Freitag 1965: Fig. 18]. According to Pearson et al. (2006: 68), distinct individuals of this form are confined to southeastern and central Arizona. The record from “Texas” (Bousquet and Larochelle 1993: 58) is in error; that from “Utah” is apparently based on intergrades.
1. According to Freitag (1965: 111), the type series of
This subspecies, the “Navajo Tiger Beetle”, inhabits a small area in the southern parts of the Rocky Mountains in Utah, southwestern Colorado, northern Arizona, and New Mexico [see Freitag 1965: Fig. 18; Pearson et al. 2006: Map 18].
This subspecies, also known as the “Western Tiger Beetle”, ranges from Vancouver Island to western Alberta, south to central Utah and southern California along the Mexican border [see Freitag 1965: Fig. 18]; also recorded from Baja California (Murray 1979: 50) and northern Sonora (Cazier 1954: 242). The records from New Mexico (Fall and Cockerell 1907: 155), “Arizona,” and “New Mexico” (Boyd 1982: 6) probably refer to the
This subspecies, the “Nova Scotia Tiger Beetle”, is known from Nova Scotia, including Cape Breton Island, Prince Edward Island, and the Magdalen Islands in Quebec (Leonard and Bell 1999: 101). The record from “New Brunswick” (Boyd 1982: 6) needs confirmation.
This subspecies, also known as the “Bronze Tiger Beetle”, ranges from Newfoundland to southwestern British Columbia, south to southern Oregon, northeastern Arizona, central Texas, and northern Florida (Choate 2003: Map 18) [see Pearson et al. 2006: Map 16]. The record from “Northwest Territories” (Boyd 1982: 6) needs confirmation.
Intergrade populations between this subspecies and the
This subspecies, the “Tanner’s Tiger Beetle”, is known only from the Green River Valley in eastern Utah (Pearson et al. 2006: 65).
This species, also known as the “Colorado Dune Tiger Beetle”, is restricted to the Great Sand Dunes National Monument and adjacent areas in Alamosa, Costilla, and Saguache Counties in south-central Colorado (Kippenhan 1990: 310; Pearson et al. 2006: 79). The record from “New Mexico” (Boyd 1982: 7) is in error.
This species, also known as the “Bruneau Dune Tiger Beetle”, has yet been found only at the type locality in southwestern Idaho [see Leffler 2001: Fig. 1].
This subfamily contains a single tribe.
This tribe includes a single genus.
Thirteen species in the Nearctic (three species, one of them Holarctic), Neotropical (two species in mountains of Middle America), and Palaearctic (nine species) Regions arrayed in three subgenera:
Ball and Erwin (1969: 883) published a key to all species then known, including the three found in North America. Lindroth (1961a: 121-125) covered the Nearctic species.
Eleven species in North America (three species), Middle America (two species), Asia (seven species), and Europe (one species which is also found in Asia and North America).
This species ranges from Kodiak Island and the Kenai Peninsula in southern Alaska south to northern California (Lindroth 1961a: 122; Humboldt County, MCZ).
This species is found along the coastal region from Washington (Hatch 1953: 64) to at least the San Francisco Bay area (LeConte 1851: 180).
This Holarctic subspecies is found on the Kuril Islands and on the Aleutian Islands and Kenai Peninsula in Alaska (Lindroth 1961a: 125).
This Holarctic subspecies ranges over most of Europe and a large part of Asia (see Bousquet 2003a: 98) and from the west coast of Alaska above the arctic circle (Lindroth 1961a: 125) to Newfoundland (Lindroth 1955a: 33), south to northern West Virginia (Tucker and Preston Counties, CMNH), western Nebraska (Kimball County, USNM), northern New Mexico (Sandoval and San Miguel Counties, UASM), southern Arizona (Greenlee County, UASM), and the San Bernardino Mountains in southern California (Fall 1901a: 40, as
This subfamily includes a single tribe.
Northern Hemisphere, with about 50 species arrayed in three genera, all represented in North America.
Three species in the arctic and subarctic areas of the Nearctic (two Holarctic species) and Palaearctic (three species) Regions.
Lindroth (1954a) reviewed the species and provided a key for their identification. The two species found in North America were also covered in his monograph of the
Goulet (1983: 447) regarded
This Holarctic subspecies is known in Asia from Kazakhstan, Mongolia, and eastern Siberia (Goulet 2003: 206) and in the Nearctic Region from a few localities in Alaska, Northwest Territories (Lindroth 1961a: 102), northern Alberta (Bourassa and Wood 2011: 144; Fort McMurray area, Gerald J. Hilchie pers. comm. 2009), and Labrador (Lindroth 1961a: 102). Fossil remnants, dated between 10,400 and 17,000 years B.P., have been unearthed in Cape Breton Island, Nova Scotia (Miller 1997: 250) and Iowa (Schwert 1992: 76).
The nominotypical subspecies is found in northern Europe and eastern Siberia.
This Holarctic species is known from scattered localities in Norway, European Russia, Siberia, Mongolia (Goulet 2003: 206), and northeastern China (Li 1992: 30, as
Eight species in the arctic, subarctic, boreal, and temperate areas of North America (six species) and Eurasia (four species). Two species-group taxa are Holarctic (
Lindroth (1954a) reviewed the species and subsequently treated all the North American species in his monograph of the
This species is found in northern European Russia and eastern Siberia in the Palaearctic Region (Goulet 2003: 206) and from western Alaska to the western shore of the Hudson Bay in northern Manitoba [see Nielsen et al. 1987: Fig. 17b; Morgan et al. 1986: Fig. 1]. Fossil remnants from the Pliocene or early Pleistocene have been unearthed in northwestern Greenland, Meighen Island, Ellesmere Island, and eastern Siberia (Böcher 1995: 20).
This Nearctic species ranges from Newfoundland (Lindroth 1955a: 31) to westernmost Yukon Territory (Goulet et al. 2009: 33), south to southern British Columbia (Lindroth 1961a: 106), northwestern Montana (Russell 1968: 46; LaBonte and Johnson 1989: 170), northern Illinois (Blatchley 1910: 50), and Massachusetts (Middlesex County, MCZ, USNM) [see Morgan et al. 1986: Fig. 4]. The record from “Pennsylvania” (Bousquet and Larochelle 1993: 86, as
This species has passed under the name
The range of this species extends from Newfoundland (Lindroth 1955a: 30) to the foothills of the Rocky Mountains in Alberta (Lindroth 1961a: 107-108), south to northern New York and New England (Lindroth 1961a: 107) [see Morgan et al. 1986: Fig. 3]. The record from Michigan (Bousquet and Larochelle 1993: 86), based on a specimen labeled from Lake Superior in CMNH, needs confirmation.
This Holarctic subspecies is known from Hokkaidō in Japan and the Russian Far Eastern Region, including Sakhalin and Kamchatka, in the Palaearctic Region and from Alaska, as far south as Anchorage, and northwestern Northwest Territories in the Nearctic Region (Goulet et al. 2009: 33).
The nominotypical subspecies ranges from the Atlantic Coast in Europe to the Lake Baikal region in Siberia (Goulet et al. 2009: 33).
This species is found mainly west of the Cascade Range from southwestern British Columbia, including Vancouver Island, to south-central Oregon (LaBonte and Johnson 1989: 171).
The range of this species extends from Newfoundland (Lindroth 1955a: 31-32) to southern Alaska, south to central British Columbia (Lindroth 1961a: 106), northwestern Montana (LaBonte and Johnson 1989: 170), northern Indiana (Blatchley 1910: 50), and New Jersey (Smith 1890: 73) [see Morgan et al. 1986: Fig. 2; Ball and Currie 1997: Fig. 3].
Thirty-nine species in the arctic, subarctic, boreal, and temperate areas of North America (19 species) and Eurasia (24 species). These species are arrayed in five subgenera, all but
Goulet (1983) revised the species then known and provided a key for their identification.
One species in the subarctic regions of North America, Asia, and Europe.
Goulet (1983) concluded from his phylogenetic analysis using adult and larval characters that this subgenus was the most basal lineage of
This Holarctic subspecies is widely distributed in the subarctic areas. In the Palaearctic Region, it ranges from the British Isles to the Kamchatka Peninsula, and in the Nearctic Region from Alaska to Labrador [see Goulet 1983: Fig. 157]. Fossil remnants of this species, dated between 10,100 and 20,700 years B.P., have been unearthed in southern Quebec (Mott et al. 1981: 146), southern Ontario (Morgan and Morgan 1981: 1107), Illinois (Schwert 1992: 76), and southeastern Iowa (Baker et al. 1986: 96); others, believed to be 2.0-2.5 million years old, have also been found in Greenland and Meighen Island (Bennike and Böcher 1990: 336; Böcher 1995: 22).
This subspecies is known only from a few localities on Kodiak Island, Alaska (Goulet 1983: 244).
Fourteen species in the Nearctic (six species) and Palaearctic (eight species) Regions.
Goulet’s (1983) phylogenetic analysis based on adult and larval characters suggests that
Fossil remnants of
This species extends throughout the temperate and boreal regions of North America from Newfoundland to Alaska, south to northern California along the west coast, to the White Mountains in eastern Arizona and to central New Mexico along the Rocky Mountains, to west-central Nebraska (Arthur County, Foster F. Purrington pers. comm. 2010), and to New Jersey along the east coast [see Goulet 1983: Fig. 160]. The record from east-central Missouri (Summers 1873: 132) needs confirmation.
This species is known from western Nevada and California, from the Oregon border to the Los Angeles area [see Goulet 1983: Fig. 161].
This species ranges from the west coast of Newfoundland to southern British Columbia, south to southeastern Utah, central Colorado, central Illinois, and New Jersey [see Goulet 1983: Fig. 161]. The species is known from only one locality west of the Rocky Mountains, in southern British Columbia.
This species ranges from Maine, southern Quebec, and Michigan south to northern Mississippi (Pontotoc County, Drew A. Hildebrandt pers. comm. 2009) and North Carolina (Northampton County, CNC) [see Goulet 1983: Fig. 159]. The record from Georgia (Fattig 1949: 11) needs confirmation; those from eastern Iowa (Wickham 1911b: 5; King 1914: 320) are probably in error.
This species ranges from Quebec to southern Manitoba, south to Nebraska and Maryland [see Goulet 1983: Fig. 158]. It is also known from one unexpected locality in central Alberta (Bousquet 1987a: 112), which suggests that the species is more widely distributed at least in the northern part of its range.
This species is known from two areas, Maryland and southeastern Virginia (Surry County, CNC), and southern Indiana, southern Illinois, and southern Tennessee (Marion County, CMNH) [see Goulet 1983: Fig. 159]. The apparent gap is probably due to inadequate samplings.
Eighteen species in North America (ten species) and Eurasia (ten species). Two species (
This subspecies ranges from Newfoundland to southwestern Alaska, south to central British Columbia, southeastern Alberta, northern Minnesota, north-central Ohio (Purrington and Stanton 1996: 44), northern New York, and Maine [see Goulet and Baum 1982: Fig. 1]. The records from “California,” “Wyoming,” “South Dakota,” “Iowa” (see Bousquet and Larochelle 1993: 87), and Missouri (Anonymous 2007) are likely based on mislabeled specimens or are in error.
This subspecies ranges from the Queen Charlotte Islands (Kavanaugh 1992: 58) to southwestern Alberta, south to central Colorado, central Idaho, and southern Oregon (Goulet and Baum 1982: 2272; Fig. 1).
This species is widely distributed from central Alaska to Nova Scotia, south to northern Virginia, central Missouri, northern New Mexico, and southern California near the Mexican border [see Goulet 1983: Fig. 172]. The records from “North Carolina,” northeastern Florida, “Louisiana,” and “Texas” [see Goulet 1983: Fig. 172] need confirmation.
This species extends from western Montana to southern Oregon, south to southernmost California, central Arizona, and southern Colorado [see Goulet 1983: Fig. 170].
This species ranges from northwestern Quebec along the James Bay (Chisasibi, Serge Laplante pers. comm. 2011) to central British Columbia, north to the Great Slave Lake in Northwest Territories, south to southern California, southern Arizona, and central Kansas [see Goulet 1983: Fig. 169]. Fossil remnants from a Plio-Pleistocene sequence have been unearthed in northwestern Greenland (Böcher 1995: 23).
This species is known from a few localities in southeastern Washington, northern California, southeastern Wyoming, and Colorado [see Goulet 1983: Fig. 168].
This species is known only from the original two specimens collected at the type locality.
This species ranges from southern Quebec to northern Minnesota, south to eastern Texas (Casey, 1924: 17, as
This species is found in the montane regions of northern Mongolia and neighbouring Russia west and south of Lake Baikal, in northeastern Siberia, and in the arctic regions from the Commander Islands in the Bering Sea to the eastern shore of James Bay in Quebec (Morgan and Pilny 1997: 146) [see Goulet 1983: Fig. 171]. Fossil remnants of this species, dated between 10,100 and 11,050 years old, have been found in southern Quebec (Mott et al. 1981: 146); others, older than 33,000 years B.P., has been unearthed in southwestern Ontario (Warner et al. 1988: 37).
According to Shilenkov (in Kryzhanovskij et al. 1995: 61), the type material of
This Holarctic species ranges from northern Scandinavia to eastern Siberia, and from Alaska to the Mackenzie River in the Northwest Territories [see Goulet 1983: Fig. 171]. Fossil remnants of this species, believed to be 2.0-2.5 million years old, have been found in Greenland (Bennike and Böcher 1990: 336; Böcher 1995: 23).
This species is known from a small area in Solano County, California [see Goulet 1983: Fig. 168].
This species, also known under the vernacular name “Delta Green Ground Beetle”, is considered an endangered species by the World Wildlife Fund and listed on the IUCN Red List of Threatened Species.
Five species in northern North America (two species) and Eurasia (four species). One species (
This Holarctic subspecies ranges from the Lena River in eastern Siberia to the Bering Sea Coast, and from Alaska to the Mackenzie River in Northwest Territories [see Goulet 1983: Fig. 173]. Fossil remnants from a Plio-Pleistocene sequence have been unearthed in northwestern Greenland (Böcher 1995: 23).
The subspecies
This species ranges from central Alaska to the Mackenzie River in Northwest Territories, south to southeastern Alberta, central Idaho, and central California along the west coast [see Goulet 1983: Fig. 173].
This subfamily includes a single tribe.
This tribe includes a single genus.
About 70 species in the Nearctic (11 species), Neotropical (six species in Middle America), Oriental (16 species), Palaearctic (16 species), and Afrotropical (20 species) Regions. The species are arrayed in two subgenera:
About 60 species in the Nearctic (11 species), Neotropical (six species in Middle America), Oriental (16 species), Palaearctic (15 species), and Afrotropical (13 species) Regions.
Benschoter and Cook (1956) revised the North American species and provided a key for their identification. Lindroth’s (1961a: 10) key covered seven species.
This species ranges from Newfoundland (Lindroth 1955a: 150) to the foothills of the Rocky Mountains in Alberta (Lindroth 1961a: 12), south to northeastern Arizona, the state of Coahuila in Mexico (Benschoter and Cook 1956: 427), and the Florida Panhandle (Peck and Thomas 1998: 15). The record from “Vera Cruz” (Benschoter and Cook 1956: 428) needs confirmation.
This species seems to be confined to the southern half of California (Benschoter and Cook 1956: 422) and the Baja California Peninsula (Erwin 2007a: 64). Old specimens simply labeled from Arizona and Texas are known (Benschoter and Cook 1956: 422) but are probably mislabeled.
This species ranges from southern California to western Colorado (Benschoter and Cook 1956: 416) and northern New Mexico (Taos County, UASM), south to Sonora, Mexico (Erwin 2007a: 65). Old specimens simply labeled from Texas are known (Benschoter and Cook 1956: 416).
This species ranges from western Wisconsin (Messer 2010: 34) to southern Nebraska (Adams County, Foster F. Purrington pers. comm. 2010), south at least to northeastern Texas (Benschoter and Cook 1956: 425), east-central Louisiana (West Feliciana Parish, Igor M. Sokolov pers. comm. 2009), and southwestern Mississippi (Lago and Zucarro 1984: 118; Wilkinson County, UASM).
This species is found along the Atlantic and Gulf of Mexico coasts, from Sable Island off the coast of Nova Scotia (Lindroth 1969a: 1108) to southern Florida (Peck and Thomas 1998: 15), west to southeastern Texas (Benschoter and Cook 1956: 420).
This species ranges from northern Nebraska to northwestern Indiana, north to the Minneapolis region in western Minnesota, south to Alabama and southern Texas (Benschoter and Cook 1956: 420, 422). The records from Wisconsin (Rauterberg 1885: 11) and Charity Island in Michigan (Andrews 1916: 72) are probably based on misidentified
This species ranges from southern California to western Texas, north to southern Utah (Benschoter and Cook 1956: 416), south to Zacatecas in Mexico (Erwin 2007a: 67). The record from “Montana” (Bousquet and Larochelle 1993: 90) is likely in error.
The range of this species extends from southwestern Saskatchewan (Ronald R. Hooper pers. comm. 1990) to Vancouver Island (Lindroth 1961a: 13), south to northern California and southern Wyoming (Benschoter and Cook 1956: 426).
This species ranges from the southern part of the Ontario Peninsula to south-central North Dakota (Benschoter and Cook 1956: 418), south to northwestern Texas (Hutchinson County, Robert L. Davidson pers. comm. 2012) and northwestern Tennessee (Lake County, CMNH); seemingly isolated in southeastern Alberta (Lindroth 1961a: 11).
This species is known from southwestern Oregon and northern California (Benschoter and Cook 1956: 425).
This species ranges from Cape Breton Island to southern Alberta, south to southwestern Arizona, northwestern Oklahoma (Cimarron County, CNC), southwestern Arkansas (Hempstead County, MCZ), and Virginia (Benschoter and Cook 1956: 424). One specimen labeled from Alameda County in western California, seen by Benschoter and Cook (1956: 424), is possibly mislabeled. The record from “Texas” (Bousquet and Larochelle 1993: 90) needs confirmation.
Worldwide, with about 1,870 species arrayed in eight tribes:
Thirty-five species arrayed in two genera:
Western Hemisphere, with 32 species arrayed in two subgenera:
Bänninger (1950) reviewed all the species and provided a key for their identification. Purrington and Drake (2005: 254-255) published a key to the North American species. A modern taxonomic revision of the genus is needed as challenging problems remain to be resolved (Ball and Bousquet 2000: 76).
Nineteen species in the temperate, subtropical, and tropical areas of North America (11 species) and Middle America (13 species).
Snow (1907: 141) recorded
This species ranges from “Utah” (LeConte 1874a: 273) to southeastern Nebraska (Pawnee County, Peter W. Messer pers. comm. 2008), south at least to eastern Arkansas (Cook and Holt 2006: 2313) and Michoacán in Mexico (Ball and Shpeley 1992a: 46).
This species ranges from southeastern New York (Notman 1928: 211) and New Jersey (Bänninger 1950: 491) to Wisconsin (Messer 2010: 34), south to southeastern Texas (Galveston County, MCZ; Bänninger 1950: 491) and central Florida (Lake County, MCZ).
The range of this species extends from northern Arizona to “Missouri” (Bänninger 1950: 492), north to northern Nebraska (Cherry and Sheridan Counties, Peter W. Messer pers. comm. 2004), south to southern Texas (Johnson 1978: 67) and northeastern Mexico (Bänninger 1950: 492).
Bänninger (1950: 510) retained
This species ranges from the southern part of the Prairie Provinces (Lindroth 1961a: 131) south to northern Sonora (Bates 1884: 264), western and northern Texas (Wheeler, Grayson, Presidio, and Brewster Counties, MCZ, CMNH; Bänninger 1950: 490), and southeastern Louisiana (Summers 1874a: 79), east to Indiana (Blatchley 1910: 55; Bänninger 1950: 490) and southwestern Michigan (Dunn 1982a: 206). The record from southwestern Virginia (Horn 1869a: 123) is probably in error.
This species is found from “Iowa” (Bänninger 1950: 492) to southeastern Wyoming (Laramie County, CMNH), south to southeastern Arizona (Casey 1913: 89, as
Bänninger (1950: 510) retained
This species ranges from New Jersey (Smith 1890: 74; Smith 1910: 201) and north-central Virginia (Carrington 2002: 107) to west-central Kansas (Snow 1878: 63; Knaus 1907: 233), south to central Texas (LeConte 1846a: 146; Lee County, MCZ) and the Florida Panhandle (Peck and Thomas 1998: 16); also recorded from Durango in Mexico (García 2004: 289, as
This species ranges from southern Arizona (Schaeffer 1905: 142) south at least to Durango in Mexico (Bänninger 1950: 493).
Bänninger (1950: 510) retained
This species ranges from southern Maryland (Peter W. Messer pers. comm. 2010) to the Florida Keys (Nichols 1988b: Fig. 5-4; Peck and Thomas 1998: 16), west to “Texas” (Leng 1915: 565; Bänninger 1950: 488), north along the Mississippi River drainage to southwestern Kentucky (Mammoth Cave National Park, CMNH).
Bänninger (1950: 509) retained
This species is probably restricted to the Coastal Plain ranging from South Carolina (Kirk 1970: 9; Ciegler 2000: 38) to southern Florida including the Keys (Peck and Thomas 1998: 17), west to southeastern Louisiana (Allen 1965: 62).
Bänninger (1950: 510) retained
This species is known from southeastern Georgia (Fattig 1949: 12), the Florida Peninsula and Panhandle (Peck and Thomas 1998: 17), and southwestern Alabama (Löding 1945: 12).
The range of this species extends from Massachusetts (Miliotis 1974: 114) to central Iowa (Purrington and Drake 2005: 256), south to southeastern Mississippi (Casey 1913: 76, as
Bänninger (1950: 510) retained
Worldwide, with about 495 species (Lorenz 2005: 135-141 and the genus
Worldwide, with about 445 species arrayed in 37 genera. The tribe is much more diversified in term of species (about 51% of the world fauna) in the Afrotropical (particularly on Madagascar) than anywhere else. The Western Hemisphere has only four endemic genus-group taxa:
About 190 species (Lorenz 2005: 137-140) arrayed in four subgenera:
Bänninger (1938) reviewed the species and provided keys for their identification. Three new North American species were subsequently described. Bousquet and Skelley (2010) published a key to all Nearctic species except
About 130 species in the Nearctic (seven species), Neotropical (about 25 species), Oriental (about 45 species), Palaearctic (about 25 species, most of them in Asia), and Afrotropical (about 45 species) Regions. The bulk of the species (about 77% of the world fauna) are found in the tropics of the Old World.
The taxonomy of the species of the
This species inhabits the Great Plains from south-central Kansas (Sedgwick County, MCZ) to the Rio Grande in Texas (Hidalgo and Cameron Counties, MCZ, USNM), west to western Texas (El Paso County, USNM), including southeastern and central New Mexico (Chaves County, CMNH; Ellis et al. 2001: 16), east to northeastern Louisiana (Franklin and Tensas Parishes, Igor M. Sokolov pers. comm. 2009). The records from “Wisconsin” and “Iowa” (Bousquet and Larochelle 1993: 96) are likely in error.
Bänninger (1938: 152) regarded this taxon as a subspecies of
This species ranges along the Coastal Plain from New Jersey (Smith 1890: 74, as
This species ranges from southern Ontario (CNC) to eastern North Dakota (Tinerella 2003: 635 as
This species is known from coastal Florida, including the Keys (Peck and Thomas 1998: 17), the Bahamas, Cuba, and the Yucatán Peninsula in southern Mexico [see Nichols 1986: Fig. 9]. One old specimen simply labeled from Louisiana is known (Nichols 1986: 261).
This species is endemic to the Florida Peninsula north of Lake Okeechobee [see Nichols 1986: Fig. 10].
This species is known only from the holotype.
This species is found from southeastern New Hampshire (Rockingham and Strafford Counties, Donald S. Chandler pers. comm. 2008) to eastern North Dakota (Cass County, Donald P. Schwert pers. comm. 1989), including southern Ontario (Lindroth 1961a: 129), south to the Yucatán Peninsula (Nichols 1988a: 52), the Florida Keys (Peck and Thomas 1998: 17), and Cuba (Darlington 1934: 67; Nichols 1988b: Fig. 5-8), west along the southwest to southwestern California (LeConte, 1852a: 198, as
Bänninger (1938: 151) retained
Worldwide, with about 820 species (Lorenz 2005: 141-150) arrayed provisionally in three subtribes,
Worldwide, with about 705 species. The Northern Hemisphere is represented by about 170 species (roughly 24% of the world fauna) and North America by 52 species (about 7% of the world fauna).
Worldwide, with about 375 species described (Lorenz 2005: 141-145) arrayed in nine subgenera:
Western Hemisphere, with at least 30 species in the Nearctic (two species, one of them adventive) and Neotropical (at least 30 species) Regions. One species, possibly adventive, is known from the suburbs of Sydney, Australia (Baehr 2008: 23-25).
Nichols (1985b: 380) discussed the structural differences between the two species found in North America.
This subgenus as defined by Kult (1947: 31) includes the species of groups 19 (mistakenly reported as group 29 by Kult) and 24 of Putzeys (1867b: 145, 166-178). Recently Dostal (2011) listed this taxon as a distinct genus and described a new subgenus,
The range of this species extends from Connecticut (Krinsky and Oliver 2001: 44) to eastern South Dakota (Kirk and Balsbaugh 1975: 16), including southernmost Ontario (Bousquet 1987a: 119), south to southern Texas (Zapeta, Kleberg, and Gonzales Counties, CMNH; Leng 1915: 570) and southern Florida (Peck and Thomas 1998: 17), west along the south to the Colorado River drainage in San Bernardino County, California (Fall 1901a: 41); also recorded from Cuba (Dejean 1825: 415; Jacquelin du Val 1857: 15), Jamaica (Nichols 1988b: Fig. 5-14), and Mexico as far south as Oaxaca (Erwin 2011b: 169). One old specimen labeled “Mass” is known (MCZ).
This species is native to South America and is adventive in North America where it is known from southeastern United States (Nichols 1985b: 380). The first inventoried specimen collected on this continent was found in Mobile, Alabama in 1948 (Nichols 1985b: 380).
Worldwide, with over 250 species described. The number of species cannot be assessed at this time since many species included by Lorenz (2005: 141-145) in this subgenus belong to
Bousquet (1997c: 347-348) published a key to all North American species and two unnamed ones. One species (
This species is known from Levy and Gilchrist Counties in northern Florida.
This European species is adventive in North America where is it known from southern Quebec (Larochelle 1975: 78) to southwestern Ohio (Dury 1879: 162) and Connecticut (Krinsky and Oliver 2001: 44), from southern Manitoba (Pollock 1991b: 298), from British Columbia (Lindroth 1961a: 162), and from Washington (Hatch 1953: 66). The first inventoried specimen collected on this continent was found prior to 1838 (Randall 1838b: 34, as
This Palaearctic subspecies is adventive in North America where it is found in the east from Newfoundland (Lindroth 1955a: 45) to northern Minnesota (Petrice et al. 2002: 9), south to northern Pennsylvania (Bradford County, CMNH) and in the west from southwestern British Columbia (Bousquet 1987a: 119) to south-central Saskatchewan (Ronald R. Hooper pers. comm. 2002), south to northwestern Wyoming (Teton County, Foster F. Purrington pers. comm. 2010) and southern Oregon (Nelson and Reynolds 1987: 12). The first inventoried specimen collected in the east was found in 1915 in the Montreal region (Lindroth 1961a: 161) and in the west in 1937 in western Washington (Hatch 1949b: 118). The records from Georgia (Fattig 1949: 14) and Alabama (Löding 1945: 12) are probably in error; those from “Ohio” (Hamilton 1889b: 93), “Illinois,” “Indiana,” and “Idaho” (Bousquet and Larochelle 1993: 102) need confirmation.
The subspecies
The range of this species, also known under the vernacular name “slender seed-corn Beetle”, extends from the Saint Lawrence Plain in southern Quebec (Lindroth 1961a: 164) to northern Utah (Knowlton and Wood 1947: 94; Davis and Utah Counties, USNM), south to southern Texas (Hlavac 1967: 31; Johnson 1978: 67) and northern Georgia (Fattig 1949: 14; Hlavac 1967: 30). At least one specimen simply labeled from Florida is known (Hlavac 1967: 30). The record from Idaho (Anonymous 1960: 642) needs confirmation.
This species is known from the holotype collected in east-central North Carolina and six old specimens labeled “D C,” “Md,” and “Ill.” (CMNH).
This species occurs from southern British Columbia (Lindroth 1961a: 163) and northern Idaho (Hatch 1953: 66) south to central Oregon (Fall 1922b: 164; Benton County, MCZ).
Hlavac (1967: 27) believed this species was morphologically distinct enough to propose a new subgenus,
This species ranges from southern Maine (Nelson 1995: 71) to “Illinois” (Hlavac 1967: 23), including southeastern Michigan (Saint Clair and Wayne Counties, CMNH) and west-central Indiana (Montgomery County, R. Michael Brattain collection), south to north-central Texas (Knaus 1905b: 348), southeastern Louisiana (Tangipahoa Parish, USNM) and the Florida Panhandle (Peck and Thomas 1998: 17).
This species ranges from South Carolina (LeConte 1857b: 82; Ciegler 2000: 42) and northern Georgia (Fattig 1949: 14; Floyd County, MCZ) west to southwestern Texas (Dajoz 2004: 117; El Paso County, MCZ), north along the Mississippi River drainage to west-central Indiana (Tippecanoe County, CMNH) and east-central Kansas (Dickinson and Douglas Counties, MCZ); also recorded from Jalisco, Nayarit, Tamaulipas (Erwin 2011b: 183) and Veracruz in Mexico (Bates 1881: 32). The record from the District of Columbia (Ulke 1902: 6) needs confirmation.
Hlavac (1967: 19) believed this species was morphologically distinct enough to propose a new subgenus,
This species ranges from southern Missouri (Hlavac 1967: 32; Ripley County, CMNH) to central Virginia (Nelson County, USNM) and the District of Columbia (Ulke 1902: 37), including southern Ohio (Washington County, UASM; Wright and Whitehouse 1941: 70), south to the Florida Panhandle (Jackson County, CNC), southwestern Mississippi (Copiah County, MCZ), and southern Texas (Zapata, San Patricio and Dimmit Counties, CMNH, UASM).
This species is found in California from Siskiyou County (CAS) to San Diego County (Moore 1937: 5).
Four species in the temperate regions of eastern North America (three species) and caves in the Sierra de Guatemala mountains in northeastern Mexico (one species,
Ball (2001) revised the species and provided a key for their identification. One species (
In a cladistic analysis of the Western Hemisphere subgenera proposed by Ball (2001: Fig. 9), this taxon turn out as the sister-group to {
This species is known only from two specimens collected in Blount County, north-central Alabama.
This species is confined to eastern United States ranging from Connecticut (Krinsky and Oliver 2001: 46) to southeastern Kansas (Knaus 1885: 57), including eastern Iowa (Ball 2001: 147), south to “Louisiana” (LeConte 1857b: 81) and central Florida (Peck and Thomas 1998: 17).
This species is known only from eastern Oklahoma (Ball 2001: 147).
This subgenus includes 72 species in the Nearctic (5 species), Palaearctic (6 species), Oriental (18 species) and Afrotropical (43 species) Regions (see Bousquet 2009: Table 2).
There is no key for the identification of the North American species of this subgenus. Nichols (1988a: 147-153) covered two species (
Until recently, the species of this subgenus were included in the subgenus
This species ranges from Massachusetts (Norfolk County, CMNH) to southwestern Wisconsin (Grant County, CMNH), south to eastern Texas (Riley 2011; San Augustine County, CMNH) and southern Florida (Nichols 1988a: 150).
This species ranges from Nova Scotia (Lindroth 1954c: 301) to southeastern North Dakota (Tinerella 2003: 635), south to western (Dajoz 2007: 23) and southern (Gonzales County, MCZ) Texas, southern Louisiana (Hine 1906: 76, as
This species is known only from the type locality.
This species is known from Harrison County in southeastern Mississippi (Drew A. Hildebrandt pers. comm. 2010), Saint Tammany Parish in southeastern Louisiana, Trinity County in eastern Texas (Bousquet 2009: 44), and LeFlore County in eastern Oklahoma (Matthew Gimmel collection). The records from southeastern New York (Leng and Beutenmüller 1893: 135), Lancaster County in Pennsylvania (Rathvon 1869: 524), northwestern Georgia (Fattig 1949: 15), and southwestern Florida (Leng 1915: 571) are in error (see Bousquet 2006c: 3).
This species ranges from southeastern New York (Notman 1928: 213) to eastern South Dakota (Kirk and Balsbaugh 1975: 17), south to eastern Oklahoma (Latimer County, UASM), “Louisiana” (LeConte 1879a: 34), and southern Florida (Peck and Thomas 1998: 17). The record from the lower peninsula of Michigan (Hubbard and Schwarz 1878: 644) needs confirmation.
At least 33 species (see Bousquet 2009: table 1) in temperate, subtropical, and tropical areas of the Nearctic (nine species) and Neotropical (29 species) Regions. Kult’s (1947: 31) statement that
There is no revision of the North American species of this genus. The last key published, that of LeConte (1879a), included seven of the eight species recorded at the time (
This taxon is very likely monophyletic, characterized by the synapomorphic condition of the lateral bead of the pronotum uninterrupted and removed from the base. The genus
The range of this species extends from Massachusetts (Purrington 1997: 96) to eastern South Dakota (Kirk and Balsbaugh 1975: 17; French et al. 2004: 557), south to southeastern Texas (Snow 1906a: 141; Cameron County, MCZ) and southern Florida including the Keys (Peck and Thomas 1998: 17), west along the southwest to southern Arizona (Dajoz 2004: 116; Cochise and Pima Counties, UASM); also recorded from several islands of the West Indies, Mexico, Honduras, and Nicaragua (Erwin 2011b: 162). The record from “New Hampshire” (Bousquet and Larochelle 1993: 103) needs confirmation.
This species is known from Long Island, New York (Cooper 1935: 144) to east-central South Carolina (Ciegler 2000: 41) and “Georgia” (LeConte 1844: 50) and from “Louisiana” (LeConte 1879a: 34) and Cuba (Chevrolat 1863: 193, as
This species has been reported from several islands of the West Indies and several countries from southern Mexico to South America (Erwin 2011b: 171). It is also known from one specimen, the holotype of
This species occurs from southeastern New York (LeConte 1857b: 82) to southeastern South Dakota (Kirk and Balsbaugh 1975: 17), south to southern Texas (Wickham 1897: 103; Johnson 1978: 67) and northern Florida (Leon County, USNM), west along southern United States to southern California (Imperial, Riverside, and Kern Counties, CAS, USNM) and the Baja California Peninsula (Horn 1894: 307).
This species has been recorded by mistake as
This species is known from northeastern Mississippi (Tishomingo County, Drew A. Hildebrandt pers. comm. 2009), southeastern Louisiana (Putzeys 1867b: 149), and some islands of the West Indies (Nichols 1988a: 143; Peck 2009b: 5).
This species ranges from southwestern Ohio (Dury 1882: 218) to southeastern South Dakota (Kirk and Balsbaugh 1975: 17), including southern Wisconsin (Messer 2010: 35), south to east-central Texas (Riley 2011), southeastern Louisiana (Jefferson Parish, MCZ; Allen 1965: 62; LeConte 1879a: 34), southern Mississippi (Hancock County, Drew A. Hildebrandt pers. comm. 2008), central Alabama (Shelby County, CMNH), and southwestern South Carolina (Ciegler 2000: 42).
This species is known for sure only from “Texas” (Putzeys 1846: 104; MCZ). The record from “Kansas” (Bousquet and Larochelle 1993: 104) needs confirmation.
This species may be restricted to the Coastal Plain and Piedmont Plateau. It is known from Staten Island, New York (Leng 1915: 569) and New Jersey (Hamilton 1889a: 30; Leng and Beutenmüller 1893: 96) south to southern Florida (Peck and Thomas 1998: 17), west to south-central Texas (Bexar County, CMNH, USNM). The record from eastern Iowa (Wickham 1911b: 5) is probably in error. One specimen simply labeled from Tennessee (CMNH) is also known.
This species is known only from the Atlantic shore of Virginia (Hoffman et al. 2006: 18), southern North Carolina (Brunswick County, Ken Karns pers. comm. 2009), southeastern South Carolina (Charleston County, USNM), Florida as far south as Collier County (USNM), and southwestern Alabama (Van Dyke 1925: 125; Baldwin County, MCZ).
Western Hemisphere, with 75 species in temperate, subtropical, and tropical areas of the Nearctic (24 species) and Neotropical (about 60 species) Regions arrayed in three subgenera:
Whitehead (1972) revised all the North American and some of the Neotropical species.
Ten species in North America (one species) and Middle and South America (ten species).
This subspecies is found in southeastern California and Arizona south to southern Sinaloa and northern Nayarit [see Whitehead 1972: Fig. 74].
The subspecies
Fifty-seven species in the Nearctic (23 species) and Neotropical (45 species) Regions.
This species is found along the Colorado River in eastern New Mexico to the Rio Grande drainage system in central Texas and Coahuila [see Whitehead 1972: Fig. 146]; it is also recorded from southwestern Oklahoma (Kondratieff et al. 2005: 173).
This species ranges from the Okanagan Valley in southern British Columbia to northwestern South Dakota, south to the state of Mexico and southern California [see Whitehead 1972: Fig. 241].
The known range of this species extends from southern Illinois to southern California, south to Baja California Sur and Colima in Mexico [see Whitehead 1972: Fig. 240].
This species ranges west of the Rocky Mountains from Vancouver Island to western Idaho, south to southern California [see Whitehead 1972: Fig. 237].
This species is known only from a few localities in northern and central California [see Whitehead 1972: Fig. 240] and from San Bernardino County in the southeast (Dajoz 2007: 20).
This widely distributed species ranges from northern California south through Arizona and western New Mexico to Colombia, including Baja California [see Whitehead 1972: Fig. 239].
This species ranges from southern Missouri to southeastern Colorado south to Nuevo León and Tamaulipas [see Whitehead 1972: Fig. 238].
This species is widely distributed east of the Mississippi River from New Brunswick (Lindroth 1961a: 167) to eastern Illinois, south to east-central Louisiana (West Feliciana Parish, Igor M. Sokolov pers. comm. 2009), southern Mississippi (Clairborne, Covington, Pearl River, Stone, and Wilkinson Counties, Drew A. Hildebrandt pers. comm. 2008), and western North Carolina (Whitehead 1972: 267, Fig. 236). Old specimens simply labeled from “Georgia,” “South Carolina,” and “Wisconsin” are known (Whitehead 1972: 267). The record from “Alabama” (Bousquet and Larochelle 1993: 106) needs confirmation.
This species ranges from southern Arizona south to Costa Rica in Pacific drainage areas [see Whitehead 1972: Fig. 101].
This species ranges from southern Quebec (Larochelle 1975: 106) to South Dakota (Kirk and Balsbaugh 1975: 16), north to southwestern Saskatchewan (CNC), south to central Texas and northern Florida, west along southern United States to southeastern Arizona [see Whitehead 1972: Fig. 100]. Also recorded from the Bahamas (Erwin 2011b: 215).
This species is currently known only from a few specimens collected in Florida, Guatemala, and Costa Rica (Whitehead 1972: 201). Also recorded from Honduras and Panama (Erwin 2011b: 218).
Whitehead (1972: 201) stated that he was uncertain if the two Central American specimens he saw were conclusively conspecific with those from Florida.
This species is distributed from New Brunswick (Webster and Bousquet 2008: 16) to northeastern Montana, south to the Rio Grande Valley in south-central Texas, Tamaulipas in Mexico, and central Alabama [see Whitehead 1972: Fig. 206]. One old specimen simply labeled from “South Carolina” is known (Whitehead 1972: 251). Except for four apparently mislabeled “Florida” specimens, the species is unknown south of New Jersey along the Atlantic Coast.
This species is known only from a few localities in southern Arizona [see Whitehead 1972: Fig. 209].
The range of this species extends from southern Arizona to Tamaulipas, south to Costa Rica [see Whitehead 1972: Fig. 208].
This species is known from eastern Arizona and southwestern New Mexico [see Whitehead 1972: Fig. 207].
This species ranges from Arizona and western New Mexico, south to Nayarit [see Whitehead 1972: Fig. 147].
This subspecies is restricted to central and western California, from Shasta County in the north to San Diego County in the south [see Whitehead 1972: Fig. 147].
The subspecies
This species ranges from Kansas south to the Rio Grande Valley in southern Texas [see Whitehead 1972: Fig. 185]. One specimen simply labeled from “Ohio” (Whitehead 1972: 229) could be mislabeled. The record from south-central Colorado (Wickham 1902: 232) is probably in error.
This species ranges from Maine, southern Quebec, and Michigan south to Tennessee and North Carolina [see Whitehead 1972: Fig. 190]; also seen from eastern Oklahoma (Le Flore County, FFPC). The records from eastern Iowa (Wickham 1911b: 6) and Missouri (Summers 1873: 133) need confirmation; that from southwestern Colorado (Wickham 1902: 232) must be in error.
Whitehead (1972: 237) noted that four specimens from Texas, without definite locality data, may represent an isolated form of this species.
This species is known only from a few specimens collected in or near the Ozark Mountains in southern Missouri, western Arkansas, and eastern Oklahoma [see Whitehead 1972: Fig. 189].
This species is known for sure only from a few specimens collected in New York, West Virginia, and Kentucky [see Whitehead 1972: Fig. 189]; it was also reported from southwestern Ohio (Dury 1910: 66) and northeastern Georgia (Fattig 1949: 15).
This species is known from a few localities throughout much of Texas [see Whitehead 1972: Fig. 189].
This species occurs from southern Texas south to southeastern Oaxaca and northern Chiapas [see Whitehead 1972: Fig. 188].
Four species are placed in this genus: one (
Whitehead (1967) and Erwin (2011a) reviewed the species and provided keys for their identification.
This species is known from southern Florida, the Yucatán Peninsula, Jamaica (Nichols 1988b: Fig. 5-12), Puerto Rico, the Dominican Republic (Whitehead 1969: 36), the Bahamas, Cuba, Cayman Islands (Peck 2005: 29) and several islands of the Lesser Antilles (Peck 2009a: 12). The species has been recorded also from the Gulf of Biafra in Africa (Bruneau de Miré 1979) and the state of Pernambuco, Brazil (Nichols 1988b: 89).
Thirteen species in the subtropical and tropical regions of the Western Hemisphere, including the West Indies, with one species reaching southeastern North America.
Nichols (1988a: 172-182) covered the four species found in the West Indies in his thesis and provided a key for their identification.
This species has been reported from southeastern Louisiana (Colby 2002: 37), southern Florida (Darlington 1935a: 161), the Bahamas (Peck and Thomas 1998: 18), Cuba (Darlington 1934: 70), Cayman Islands and Puerto Rico (Peck 2005: 29), Dominican Republic (Erwin 2011b: 204), Jamaica, and eastern Mexico in Veracruz and Campeche (Nichols 1988a: 181).
Western Hemisphere, with about 90 species, arrayed in three genera. All but five species are found in the Neotropical Region. One of the North American species has been collected in Japan.
Bousquet (2006c) revised the North American species and provided a key for their identification.
This genus contains 44 species (Valdés 2009: 70) restricted to the temperate, subtropical, and tropical regions of the Nearctic (two species) and Neotropical (42 species) Regions, including the West Indies.
This species ranges along the Coastal Plain and Piedmont Plateau from New Jersey (Smith 1890: 76; Smith 1910: 202) and southeastern Pennsylvania (Rathvon 1969: 524) to southern Florida, west to central Louisiana [see Bousquet 2006c: Fig. 35].
This species is found from South Carolina (Ciegler 2000: 43) to southern Florida, west to the Rio Grande in south-central Texas, north along the Mississippi River drainage to southern Illinois [see Bousquet 2006c: Fig. 34].
Twenty species (Valdés 2012) restricted to the temperate, subtropical, and tropical regions of the Western Hemisphere, including the West Indies.
This species ranges from southeastern Iowa (Wickham 1911b: 6; King 1914: 321) to western Kentucky, south to the Rio Grande in southeastern Texas, east to southern Florida, north along the coast to southern Virginia [see Bousquet 2006c: Fig. 33]. The record from southeastern Pennsylvania (Rathvon 1869: 524) is probably in error.
The range of this species extends from Long Island, New York to southwestern Wisconsin (Messer 2010: 35), south to the Rio Grande in south-central Texas and southern Florida [see Bousquet 2006c: Fig. 32]; also recorded from the Bahamas (Turnbow and Thomas 2008: 15).
Twenty-six species (Lorenz 2005: 146-147) in the temperate, subtropical, and tropical regions of the Western Hemisphere, including the West Indies. One species only is found in North America.
This species ranges over eastern United States from Washington D.C. to northeastern Kansas, including southeastern Iowa (Wickham 1911b: 6; King 1914: 323), south to the big bend along the Rio Grande in Texas and northeastern Mexico (Bousquet 2006c: 9) and southern Florida; also known from southeastern Arizona [see Bousquet 2006c: Fig. 31]. Two specimens labeled from Klamath County, Oregon (AMNH) and Gallatin County, Montana (CAS) are known (Bousquet 2006c: 9). The species has been recorded also from Japan (Habu 1963: 19).
Worldwide, with about 300 species arrayed in nine genera:
Two species restricted to the seashore of the Pacific in North America.
The two species can be differentiated using Lindroth’s (1961a) and Bousquet’s (1988a) keys to
This taxon has been considered a subgenus of
This species is confined to the seashore of the Pacific Coast of California, as far north as San Mateo County (CAS), and of the Baja California Peninsula (CNC).
This species ranges along the seashore of the Pacific from Kunghit Island in the Queen Charlotte Islands (James C. Bergdahl pers. comm. 2009) south at least to central California (LeConte 1867b: 363; San Mateo County, CAS).
About 245 species (Lorenz 2005: 151-154) in the Nearctic (about 60 species, one of them adventive), Neotropical (about 20 species, five shared with North America), Oriental (about 25 species), Palaearctic (about 140 species, many shared with the Oriental and Afrotropical Regions), and Afrotropical (about 30 species) Regions.
The North American species have never been revised. Bousquet (1988a) published a key to all species found in the area and briefly discussed each species-group. Subsequently five new species have been described by Bousquet (1997a) and Dajoz (2004), an adventive species has been detected (
Fedorenko (1996) recognized two genera within
Fedorenko (1996) recognized five subgenera within “
This species ranges from Maine (Larochelle and Larivière 1990a: 28; Androscoggin County, CNC) to southeastern Montana (Herman 1986: 62), south to westernmost (Fall 1901a: 209, as
The range of this species extends from eastern Washington (Hatch 1953: 67) to southern California (Fall 1901a: 41; Moore 1937: 5), east to southwestern Louisiana (Cameron Parish, LSAM), south to Veracruz in Mexico (Kult 1950a: 138, as
This species is found from southern New Hampshire (Hillsborough County, Ross T. Bell pers. comm. 2008) to “North Dakota” (Donald P. Schwert pers. comm. 1989), south to eastern Texas (San Augustine and Wood Counties, CMNH; Bousquet 1988a: 373; Riley 2011) and southern Florida (Peck and Thomas 1998: 17). The record from “New Mexico” (Bousquet and Larochelle 1993: 99) needs confirmation.
This species ranges from Massachusetts (Hampshire County, CNC) to northeastern Kansas (Douglas County, MCZ), including central Wisconsin (Messer 2010: 34) and southeastern Iowa (Des Moines County, MCZ), south to San Luis Potosí in Mexico (Bousquet 1988a: 373) and the Florida Panhandle (Peck and Thomas 1998: 17), west along southern United States to southeastern California (Imperial County, CAS; Dajoz 2007: 16). One old specimen simply labeled “Neb” is known (MCZ).
This species is known from scattered localities from the Saint Lawrence Plain in southern Quebec (Larochelle 1975: 81; Bousquet 1987a: 112) to southeastern Michigan (LeConte 1878c: 593), south to southeastern Pennsylvania (Dauphin County, MCZ; Lindroth 1961a: 139); also known from northeastern Nebraska (Clopton 1991: 61).
This species is known only from a few specimens collected in southern Alberta (Lindroth 1961a: 139; Bousquet 1987a: 113), northeastern Colorado (Bell 1971: 56; Lavigne 1978: 104), and northeastern Arizona (Apache County, UASM).
This species is known only from the holotype collected in south-central Manitoba.
This species is known from the type locality in southeastern Texas, from Jackson County in southeastern Mississippi (CMNH), and from two localities in the Florida Panhandle (Frank 1985: 481; Herman 1986: 63).
This species is known only from the holotype collected along the Gulf Coast of Florida.
This species is known only from a few localities along the Gulf Coast in southern Texas (Bousquet 1988a: 371; FMNH).
This species ranges from southern Labrador to the Kenai Peninsula in Alaska (Lindroth 1961a: 152, as
This species is known from a few montane locations in south-central British Columbia (Lindroth 1961a: 152; Bousquet 1987a: 116), Idaho (Bear Lake County, CNC), and the Sierra Nevada of California (Papp 1978: 165; Mono and Placer Counties, USNM). The record from “Wyoming” (Bousquet and Larochelle 1993: 101) needs confirmation.
This species is known only from two specimens collected at the type locality in southeastern Arizona.
This species is known from a few montane locations from southwestern Alberta to south-central British Columbia (Lindroth 1961a: 152; Bousquet 1987a: 116).
This species ranges from southwestern Washington (Herman 1986: 60) to Riverside County (CAS) in southeastern California and “the middle Sierras” (Fall 1901a: 41). The records from southeastern British Columbia and northern Idaho (Hatch 1953: 67) are probably in error.
This species ranges from Newfoundland (Lindroth 1955a: 42, as
This species ranges from southwestern Oregon (Herman 1986: 60) to southern California (Fall 1901a: 41; San Diego County, CAS). The record from “Arizona” (Bousquet and Larochelle 1993: 101) needs confirmation.
This European species is adventive in North America where it is known only from the Vancouver area in southwestern British Columbia (Bousquet 1997a: 94, as
The range of this species extends from Newfoundland to central Alaska, south to southern British Columbia (Lindroth 1961a: 155), northern Arizona (Coconino County, CMNH), southern New Mexico (Otero County, CNC), east-central Texas (Riley 2011), southeastern Louisiana (Summers 1874a: 79), northern Mississippi (Drew A. Hildebrandt pers. comm. 2007), and southern Florida (Peck and Thomas 1998: 17).
This species ranges from Labrador and the Ungava Bay area to Alaska, south to south-central British Columbia; isolated on the Shickshock Mountains in Gaspé Peninsula, Quebec [see Bousquet 1987a: map 1]. Fossil remnants, dated between 10,400 and 12,600 years B.P., have been unearthed in Cape Breton Island, Nova Scotia (Miller 1997: 250).
This taxon has been treated as a subspecies of
This species is found from Newfoundland (Lindroth 1955a: 43) to south-central British Columbia (Lindroth 1961a: 156), south to northwestern Minnesota (Gandhi et al. 2005: 924). The records from South Dakota (Kirk and Balsbaugh 1975: 16), Indiana (Blatchley 1910: 57), the upper and lower peninsulas of Michigan (Hubbard and Schwarz 1878: 627, 644; Hatch 1925: 548; Silvey 1936: 657), Ohio (Leng and Beutenmüller 1893: 91; Dury 1902: 110), and “Pennsylvania” (Bousquet and Larochelle 1993: 101) are in error.
This Holarctic species ranges from Scandinavia (Balkenohl 2003: 226) to the Hudson Bay in northeastern Manitoba (Garry 1993: 94, as
This species has long been known under the name
This species is known from the southern part of the Prairie Provinces (Lindroth 1961a: 155; Bousquet 1987a: 118) and from northwestern Montana (Teton County, CNC). The records from British Columbia (Jarrett and Scudder 2001: 382), “Wyoming,” and “Minnesota” (Bousquet and Larochelle 1993: 101) need confirmation.
This subspecies is found from Alaska to the Great Slave Lake area in Northwest Territories (Lindroth 1961a: 151).
The subspecies
This species is known only from southern Manitoba (Lindroth 1961a: 154) and Alberta (Langor et al. 2006: 13).
This species is known only from the type locality in southwestern North Carolina.
This species is found from the foothills of the Rocky Mountains in southern Alberta to south-central British Columbia (Lindroth 1961a: 140), south at least to central Oregon (Benton and Lincoln Counties, CNC, MCZ, USNM). One old specimen labeled “Cal” is known (MCZ).
This species is confined to the Coastal Plain ranging from North Carolina to central Florida, west to southeastern Louisiana (Tangipahoa and Saint Tammany Parishes, LSAM) [see Bousquet 1988a: Fig. 38]. One specimen simply labeled from Texas (CMNH) is known.
This species ranges from southern New Brunswick (Webster and Bousquet 2008: 16) to southern Manitoba, south to eastern Texas, southeastern Louisiana (LeConte 1857b: 80; East Baton Rouge Parish, LSAM), Mississippi (Bolivar and Issaquena Counties, Drew A. Hildebrandt pers. comm. 2009), and northeastern North Carolina [see Bousquet 1988a: Fig. 38]. The records from southwestern Georgia (Fattig 1949: 13) and southwestern Alabama (Löding 1945: 12) probably refer to
This species is found from Prince Edward Island (Larochelle and Larivière 1990a: 27) to south-central British Columbia (Lindroth 1961a: 157), south to eastern Washington (Hatch 1953: 68, as
This species is known only from the holotype collected in northeastern Kansas.
This species is known from southern Manitoba (Lindroth 1961a: 148), North Dakota (Grand Forks County, CNC, UASM), southeastern Nebraska (Foster F. Purrington pers. comm. 2010), Kansas (Stafford County, CNC, UASM), New Mexico (Chaves County, CNC, UASM), northwestern Texas (Hutchinson County, Darren A. Pollock pers. comm. 2011), and southern Florida (Monroe County, FFPC, UASM).
This species is known only from north-central Oklahoma (Herman 1986: 61), southeastern Texas (Putzeys 1846: 52; Larson 1968: 1110, as
This species ranges from southern New Brunswick (Webster and Bousquet 2008: 16) to southeastern South Dakota (Kirk and Balsbaugh 1975: 16), south to southwestern Oklahoma (Kondratieff et al. 2005: 172), southern Louisiana (Saint Martin Parish, LSAM), northern Mississippi (Bolivar, Marshall, and Warren Counties, Peter W. Messer pers. comm. 2009), and Delaware [see Bousquet 1988a: Fig. 37]. The records from the Bahamas (Turnbow and Thomas 2008: 12) and Horn Island, Mississippi (Richmond 1968: 233) refer to
Bulirsch (2009: 19) reported that
This species is found along the Atlantic Coast from western Newfoundland (Lindroth 1955a: 45, as
This species is confined to the Pacific Coast ranging from the Queen Charlotte Islands (Kavanaugh 1992: 59) south at least to northern California (Mendocino County, CNC).
This rarely collected species is known from scattered localities in the southern parts of the Prairie Provinces (Lindroth 1961a: 147; Bousquet 1988a: 373), southeastern Oregon (Harney County, CNC), northwestern (Pershing County, MCZ) and west-central (Bechtel et al. 1983: 474) Nevada, north-central Utah (Utah County, USNM), and eastern California (Fall 1925: 310, as
This Holarctic subspecies is known from the British Islands to eastern Siberia, as far south as Italy, Bulgaria, Iran, Turkmenistan, Kyrgyzstan, and Mongolia (Balkenohl 2003: 226), and in the Nearctic Region from Alaska (Lindroth 1961a: 146) to Newfoundland (Lindroth 1955a: 36), south to southwestern Pennsylvania (Allegheny County, CMNH), eastern South Dakota (Kirk and Balsbaugh 1975: 16), northern New Mexico (Rio Arriba County, USNM), northeastern Nevada (Elko County, MCZ), and southwestern California (Los Angeles County, CAS).
The subspecies
The range of this species extends from Newfoundland (Lindroth 1955a: 44) to south-central British Columbia, south to eastern Oregon (Baker County, MCZ), southeastern Arizona (Greenlee and Graham Counties, CMNH), central New Mexico (Socorro County, CNC), southern Texas (Herman 1986: 61; Dajoz 2004: 117), eastern Tennessee (Knox County, MCZ), and North Carolina (Herman 1986: 63). The records from “Georgia” (J.E. LeConte 1849: 25), Florida (Leng 1915: 568), and San Bernardino County in southwestern California (Riley 1893: 239) need confirmation.
This species ranges from southern Manitoba to south-central British Columbia, north to east-central Alaska (Lindroth 1961a: 147), south to southern California (Fall 1901a: 41), central Arizona (Griffith 1900: 565), and southern Colorado (LeConte 1879d: 500; Wickham 1902: 232; Douglas County, CNC); also recorded from Baja California Sur (Horn 1895: 225). One old specimen labeled “Pittsburg VI Pa” (CMNH) is known.
This species inhabits the Coastal Plain ranging from the coast of southern North Carolina (Brunswick County, Ken Karns pers. comm. 2009) to southern Florida (Nichols 1988b: Fig. 5-5; Peck and Thomas 1998: 17), west to southeastern Texas (Putzeys 1867b: 51).
According to Whitehead (1970: 185), members of
This species ranges from west-central Wisconsin (Messer 2010: 34) and southern Manitoba (Lindroth 1961a: 142, as
This species is known from southern Maine (Kennebec County, Robert E. Nelson pers. comm. 1989), Connecticut (Litchfield County, CMNH) and “Rhode Island” (Sikes 2003: 7) in the northeast and from the Florida Panhandle (Peter W. Messer pers. comm. 2008) to southeastern Texas (Putzeys 1846: 42; Whitehead 1970: 186; Herman 1986: 61) in the southeast.
This species ranges from southern Quebec (Larochelle 1975: 82) to south-central British Columbia (Lindroth 1961a: 141), south to the Sierra Nevada in California (Dajoz 2004: 119), southeastern Arizona (Graham County, CNC), south-central Colorado (Wickham 1902: 232), Nebraska (Clopton 1991: 61), Wisconsin (Messer 2010: 34), and “Michigan” (Garry A. Dunn pers. comm. 1986). One old specimen simply labeled from New Mexico is known (Fall and Cockerell 1907: 156).
This species is known only from the two specimens collected at the type locality in eastern California.
This species is known along the Coastal Plain and Piedmont Plateau from southeastern New York (Notman 1928: 212) to southern Florida including the Keys (Peck and Thomas 1998: 17), west to southern Texas (Aransas and Brooks Counties, CNC), and also from Minnesota (Gandhi et al. 2005: 924) and west-central Wisconsin (Messer 2010: 34). The record from southern Arizona (Wickham 1898: 300) is probably in error (Nichols 1988a: 209).
This species is known only from the type locality in southern Ontario, from Monroe and Sheboygan Counties in Wisconsin (Purrington and Maxwell 1998: 190; Messer 2010: 35), and from Highlands County in central Florida (Vince Golia collection).
This species is known only from eight specimens collected at the type locality in southeastern California.
This species is found along the Atlantic and Gulf of Mexico coasts from Maine (Larochelle and Larivière 1990a: 27, as
This species is known from the Prairie Provinces (Lindroth 1961a: 148), “Montana” (Bousquet 1988a: 372), Wyoming (Park County, CNC), and “North Dakota” (Bousquet 1988a: 372). The record from “Idaho” (Bousquet and Larochelle 1993: 98) needs confirmation.
This species ranges from the southern part of the Prairie Provinces (Lindroth 1961a: 143; Bousquet 1987a: 115) to southern Arizona (Bousquet 1987a: 115; Dajoz 2007: 21), southern Texas, and central Arkansas, east to northeastern Ohio [see Davidson and Lee 1990: Fig. 1)].
The range of this species extends from southern Quebec (Larochelle 1975: 82) to southern Alberta (Bousquet 1987a: 115), south to northern Texas (Bowie and Winkler Counties, CMNH, USNM), southeastern Louisiana (West Baton Rouge Parish, LSAM), and southern Florida (Herman 1986: 63).
This species has been reported from southern Oregon (Herman 1986: 60), western Nevada (Bousquet 1988a: 376), and Utah (Knowlton 1939: 2; Lindroth 1961a: 144; Herman 1986: 61). The records from South Dakota (Kirk and Balsbaugh 1975: 16) and Albuquerque in New Mexico (Fall and Cockerell 1907: 156) need confirmation.
This species ranges from Newfoundland (Lindroth 1955a: 45) to central Florida (Peck and Thomas 1998: 17), west to northeastern Texas (Lindroth 1961a: 143), including southeastern Mississippi (George and Greene Counties, Drew A. Hildebrandt pers. comm. 2008), and north to southern North Dakota (Burleigh County, Donald P. Schwert pers. comm. 1989) and eastern Minnesota (Gandhi et al. 2005: 924). The record from “Pennsylvania” (Bousquet and Larochelle 1993: 100) needs confirmation.
This species ranges from the southern part of the Prairie Provinces (Lindroth 1961a: 139) south to east-central California (Dajoz 2004: 119), northern Arizona (Van Dyke 1943: 22, as
This species is found along the Pacific Coast in southern California (LeConte 1852a: 196). The record from “Oregon” (Leng 1920: 47) needs confirmation; that from southwestern British Columbia (Hatch 1953: 67) is in error.
This species is found from northern Washington (Lindroth 1961a: 138) to western Montana (Russell 1968: 47), south to southwestern New Mexico (Sierra County, CMNH) and southwestern California (LeConte 1852a: 195; Moore 1937: 5). The records from “Nebraska” (Bousquet and Larochelle 1993: 98) and northwestern British Columbia (Hatch 1953: 67) are probably in error. One old specimen simply labeled “Van” is known (MCZ).
This species is found in southern California (Fall 1901a: 207).
This species ranges from “Washington” (Hatch 1953: 67) to western Montana (Jefferson County, CNC), south to northeastern Nevada (Elko County, CNC) and southern California (Fall 1910: 93; San Diego County, CAS, UASM). The record from “Wyoming” (Bousquet and Larochelle 1993: 98) needs confirmation.
Eight species in North America (two species) and South Africa (six species) arrayed in five genera:
Two species restricted to western North America.
Lindroth (1961a: 125-128) commented on the structural differences between the two species.
The status of the two forms as distinct species is questionable in my opinion. Van Dyke (1925: 123) considered the two forms as conspecific.
This species is found along the Pacific Coast from southwestern British Columbia, including Vancouver Island (Lindroth 1961a: 128), to central California (Casey 1913: 94, as
This species is known from “Oregon” south at least to central California (Lindroth 1961a: 127; Fresno County, CAS), east to Washoe County in northwestern Nevada (La Rivers 1947: 139; Lindroth 1961a: 127).
One tribe is included in this subfamily.
About 290 species (Häckel et al. 2010) in the Nearctic (four species, one of them adventive), Neotropical (about 30 species, one in Mexico, the other ones in South America), Australian (about 180 species), Oriental (five species), and Palaearctic (about 70 species) Regions placed in 34 genera. The genera are arrayed in five subtribes (Roig-Juñent 2000):
About 75 species in the Nearctic (four species), Oriental (four species), and Palaearctic (about 65 species) Regions. One species (
One Holarctic species in the subarctic and boreal regions.
Lindroth (1961a: 170-171) covered the species.
Cladistic analysis of broscine genera performed by Roig-Juñent (2000) placed this genus as the sister-group of the Holarctic genus
This circumpolar species ranges from Ireland to the Bering Sea coast (Bousquet 2003b: 237) and from Alaska (Lindroth 1961a: 171) to Newfoundland (Lindroth 1955a: 133), south to northeastern New York (Notman 1928: 241), the upper peninsula of Michigan (Chippewa County, MCZ), northern Wisconsin (Bayfield County, MCZ), northern Colorado (Dajoz 1989: 337; Gilpin County, CMNH), southern Montana, and northern Washington (Hatch 1933b: 7). Fossil remnants of this species, believed to be 2.0-2.5 million years old, have been found in Greenland (Bennike and Böcher 1990: 336; Böcher 1995: 23); others, about 20,530 years B.P., have been unearthen in northeastern Iowa (Woodman et al. 1996: 17).
Northern Hemisphere, with four species in the Nearctic (one species) and Palaearctic (three species) Regions arrayed in two subgenera:
Cladistic analysis of broscine genera performed by Roig-Juñent (2000) placed this genus as the sister-group of the Holarctic genus
One species in the temperate regions of western North America.
Lindroth (1961a: 171-172) treated the species.
This species is confined to the Pacific Coast and adjacent Coast Ranges, ranging from the Alexander Archipelago in southeastern Alaska to western Oregon (Lindroth 1961a: 172). The record from southeastern Wyoming (Lavigne 1977: 45) must be based on a mislabeled specimen.
One species in temperate western North America.
The species is covered in Lindroth’s (1961a: 172-173) monograph.
Cladistic analysis of broscine genera performed by Roig-Juñent (2000) placed this genus as the sister-group of the Asian genus
This species ranges from the southern part of the Alexander Archipelago to northwestern California, east to the Bitter Root Mountains in southwestern Montana (Ball 1956b: 34, Fig. 1).
Based on variation in color, lustre, and surface sculpture in adults, Ball (1956b) recognized a western and eastern “races” for this species.
Twenty-three species in the Palaearctic Region, one of them adventive in eastern North America.
Larochelle and Larivière (1989a) provided a description of the external structures as well as the male and female genitalia of the species found in North America.
Cladistic analysis of broscine genera performed by Roig-Juñent (2000) placed this genus as the sister-group of the temperate Asian genus
This Palaearctic species is adventive in North America where it is known from Cape Breton Island and eastern Prince Edward Island (Larochelle and Larivière 1989a: Fig. 4). The first inventoried specimen found on this continent was caught in 1987.
Five species placed in two subtribes:
Baehr et al. (2009: 106) provided a key to all species of this subfamily.
Three genera and five species are placed in this tribe.
One North American species belongs to this subtribe.
One species in western North America.
The species was covered in Lindroth’s (1961a: 4-5) monograph on the carabids of Canada and Alaska.
This minute species ranges from central Alaska (
Worldwide, with about 5,410 species arrayed in four tribes:
Worldwide, with about 2,650 species arrayed in two subtribes:
Worldwide, with about 2,470 species (Lorenz 2005: 168-200). The North American fauna is represented by about 225 species (roughly 9% of the world fauna) arrayed in nine genera.
Northern Hemisphere, with six species in temperate areas of the Nearctic (one western species) and Palaearctic (five species) Regions.
Barr (1972) provided a description of the external structures and male genitalia of the North American species.
This species is known only from a few localities in the Willamette Valley in northwestern Oregon.
About 145 described species, though more than 220 are known (Barr 2004: 1), restricted to eastern North America south of the last glaciation.
Barr (2004: 11-16) provided a key for the identification of the 26 species groups currently recognized. Barr (1959) revised the species of the
This species is known from a number of caves in DeKalb County, northeastern Alabama (Barr 2004: 40).
This species is known only from a few caves in Chattooga and Walker Counties, northwestern Georgia (Barr 2004: 40).
This species is known only from two caves in Hart and Edmonson Counties, Kentucky (Barr 2004: 24).
This species is known only from two caves in Lawrence County, southern Indiana (Barr 2004: 24).
This species is known from several caves in Carter and Elliott Counties, northeastern Kentucky (Barr 2004: 24).
This species is known from several caves in southern Clark County, southern Indiana (Barr 2004: 24).
This species is known only from two nearby caves in Jefferson County, north-central Kentucky (Barr 2004: 24).
This species is known from several caves in Rutherford and Wilson Counties, Tennessee (Barr 2004: 33).
This species is known only from the type-locality cave located two miles west of Southport in west-central Tennessee (Barr 2004: 34).
This species is known only from the type-locality cave in south-central Tennessee (Barr 2004: 33).
This species is known from a few caves in Smith and Macon Counties, northern Tennessee (Barr 1980: 91; Barr 2004: 33).
This species is known only from the type-locality cave in northern Tennessee (Barr 2004: 34).
This species is known only from the type-locality cave in north-central Tennessee (Barr 2004: 33).
This species is known from two nearby caves in Lewis and Hickman Counties, western Tennessee (Barr 1980: 93; Barr 2004: 33).
This species is known from several caves in Smith, Putnam, and Jackson Counties, northern Tennessee (Barr 1980: 92; Barr 2004: 33).
This species is still known only from two nearby caves in central Tennessee (Barr 1980: 92; Barr 2004: 33).
This species is known only from Carroll and Riley Creek caves, Coffee County, south-central Tennessee (Barr 1980: 93; Barr 2004: 33).
This species is known from a few caves in Madison County, northern Alabama (Barr 2004: 35).
This species is known only from the type-locality cave in southwestern Virginia (Barr 2004: 34).
This species is known from a few caves in Morgan County, northern Alabama (Barr 2004: 36).
This species is known only from the type-locality cave in northeastern Tennessee (Barr 2004: 34).
This species is known only from the type-locality cave in northwestern Georgia (Barr 2004: 35).
This species is known from several caves in Marshall and Morgan Counties, northern Alabama (Barr 2004: 36).
This species is known only from a number of caves in Hamilton County, southeastern Tennessee, and Dade County, northwestern Georgia (Barr 1981: 49; Barr 2004: 35).
This species is known from a few caves in Perry and Wayne Counties, Tennessee (Barr 2004: 36).
This species has been found yet only at the type-locality cave in southwestern Virginia (Barr 2004: 35).
This species is known from a few caves in Grundy and Franklin Counties, southern Tennessee (Barr 2004: 35).
This species is known from several caves in Madison County, northern Alabama (Barr 2004: 35).
This species is known from a few caves in northern Alabama (Barr 2004: 36).
This species is known only from the type-locality cave in southern Tennessee (Barr 2004: 35).
This species is known only from the type-locality cave in east-central Tennessee (Barr 2004: 35).
This species is known from a few caves in northern Alabama (Barr 2004: 35).
This species has been reported from a number of caves in Claiborne and Hancock Counties in northeastern Tennessee and Lee County in southwestern Virginia (Barr 2004: 35).
This species is known only from the type-locality cave in northeastern Alabama (Barr 2004: 35).
This species is known only from the type-locality cave in northern Tennessee (Barr 2004: 35).
This species is known from a few caves in Blount County, north-central Alabama (Barr 2004: 35).
This species is known only from the type-locality cave in eastern Tennessee (Barr 2004: 34).
This species is known only from two caves in central Kentucky (Barr 2004: 27).
This species is known from two caves in Crawford and Harrison Counties, southern Indiana (Barr 2004: 27).
This species is known from a few caves in Giles and Craig Counties, western Virginia (Barr 2004: 21).
This species is known only from the type-locality cave, near Onego, in eastern West Virginia (Barr 2004: 21).
This subspecies is known only from a few caves in Greenbrier and Monroe Counties, southeastern West Virginia (Barr 2004: 17).
This subspecies is known from several caves in Greenbrier County, southeastern West Virginia (Barr 2004: 17).
This subspecies is known only from a few caves in Greenbrier and Monroe Counties, southeastern West Virginia (Barr 2004: 17).
This subspecies is known from several caves in Greenbrier County, southeastern West Virginia (Barr 2004: 17).
This species is known from a few caves in southern Greenbrier County, West Virginia (Barr 2004: 17).
This species is known only from a few caves in southern Pocahontas County, east-central West Virginia (Barr 2004: 17).
This species is known only from a few specimens collected at the type-locality cave in northeastern West Virginia (Barr 2004: 18).
This species is known only from two caves in Tucker County, northeastern West Virginia (Barr 2004: 18).
This species is known only from a few caves in Monroe County, southeastern West Virginia (Barr 2004: 17).
This species is known only from two nearby places in the Yew Mountains, east-central West Virginia (Barr 2004: 17).
This species is still known only from the type-locality cave in southwestern Virginia (Barr 2004: 18).
This species is known only from two caves in DeKalb County, northeastern Alabama (Barr 2004: 37).
This species is known from several caves in Lee County, southwestern Virginia (Barr 1981: 60; Barr 2004: 37).
This species is known from caves in Hamilton County, Tennessee, and Dade County, Georgia (Barr 1981: 64; Barr 2004: 37).
This species is known from two nearby caves in Cumberland Gap National Park, Lee County, southwestern Virginia (Barr 1981: 60; Barr 2004: 37).
This species is known only from the type-locality cave in southeastern Tennessee (Barr 2004: 37).
This species is known only from the type-locality cave in southwestern Virginia (Barr 2004: 37).
This species is known only from the type-locality cave in southern Tennessee (Barr 2004: 37).
This species is known from a few caves in Woodford and Jessamine Counties, Kentucky (Barr 2004: 22).
This species is known only from the type-locality cave in east-central Kentucky (Barr 2004: 22).
This species is known from a few caves in Jefferson, Jennings, and Clark Counties, southern Indiana (Krekeler 1973: 51; Barr 2004: 23).
This species is known from a few widely scattered caves in Clark, Henry, Scott, and Owen Counties, Kentucky (Krekeler 1973: 49; Barr 2004: 22).
According to Barr (2004: 22), the Henry and Owen County populations possibly represent a distinct species.
This species is known from a few caves in Mercer and Garrard Counties, central Kentucky (Barr 2004: 22).
This species is known only from the holotype collected in Lee County, not Estill County as reported by Krekeler (1973: 53), in eastern Kentucky (Barr 2004: 23).
This species is known from several caves in Fayette County, north-central Kentucky (Barr 2004: 22).
This species is known only from the type-locality cave in southern Ohio (Barr 2004: 23).
This species is known only from the type-locality cave, 35 miles northeast of Lexington, northern Kentucky (Barr 2004: 22).
This species is known only from the type-locality cave in southern Ohio (Barr 2004: 23).
This species is known only from the type-locality cave in east-central Kentucky (Barr 2004: 23).
This species is known from a few caves in Woodford and Jessamine Counties, Kentucky (Barr 2004: 22).
This species is known only from the type-locality cave in northern Kentucky (Barr 2004: 22).
This species is known only from the type-locality cave in west-central Virginia (Barr 2004: 18).
This species is known only from the type-locality cave in northern Virginia (Barr 2004: 18).
This species is known only from the type-locality cave in western Virginia (Barr 2004: 18).
This species is known from a few nearby caves in Shenandoah County, northern Virginia (Barr 2004: 18).
This species is known only from the type-locality cave in northern Virginia (Barr 2004: 18).
This species is known from two nearby caves in Pendleton County, eastern West Virginia, and Highland County, western Virginia (Barr 2004: 18).
This species is known from a few nearby caves in eastern West Virginia (Barr 2004: 19).
This species is known from two caves in Giles County, western Virginia (Barr 1981: 69; Barr 2004: 38).
This species is known only from the type-locality cave in southwestern Virginia (Barr 2004: 38).
This species is known only from the type-locality cave in northeastern Tennessee (Barr 2004: 38).
This species is known only from two nearby caves near Eggleston, western Virginia (Barr 2004: 38).
This species is known only from two caves in Russell and Scott Counties, southwestern Virginia (Barr 2004: 38).
This species is known from a number of caves in Tazewell County, southwestern Virginia (Barr 1981: 70; Barr 2004: 38).
This species is known only from the type-locality cave in Whitley County, southeastern Kentucky (Barr 2004: 40).
This species is known only from the type-locality cave in southeastern Kentucky (Barr 2004: 40).
This species is known from the type-locality cave in Pike County and two abandoned coal mines in Floyd County, eastern Kentucky (Barr 2004: 40).
This species is known only from the type-locality cave in southwestern Virginia (Barr 2004: 40).
This species is known only from the type-locality cave in southeastern Kentucky (Barr 2004: 40).
This species is known only from two caves in Larue and Nelson Counties, central Kentucky (Barr 2004: 21).
This species is known from a few caves located in Mammoth Cave National Park, Kentucky (Barr 2004: 21).
This species is known from several caves in Green, Hart, and Taylor Counties, central Kentucky (Barr 2004: 21).
This species is known only from the type-locality cave in central Kentucky (Barr 2004: 21).
This species is known only from two nearby caves in Mercer and Boyle Counties, Kentucky (Barr 2004: 21).
This species is known from several caves in Garrard, Fayette, Woodford, and Owen Counties, Kentucky (Barr 2004: 21).
This species is known from several caves in Grundy and Franklin Counties, southern Tennessee (Barr 2004: 29).
This species is known only from several caves in Warren and Grundy Counties, Tennessee (Barr 2004: 29).
This species is known from several caves in Warren and Grundy Counties, Tennessee (Barr 2004: 29).
This species is known only from caves in Van Buren County, central Tennessee (Barr 2004: 29).
This species is known only from a few caves in Wise County, southwestern Virginia (Barr 2004: 39).
This species is known only from three caves in Grassy Cove, Cumberland County, in east-central Tennessee (Barr 1981: 73; Barr 2004: 39).
This species is known from two caves in Lee County, southwestern Virginia, and Hancock County, northeastern Tennessee (Barr 1981: 80; Barr 2004: 39).
This species is known from a few caves in Claiborne County, northeastern Tennessee (Barr 2004: 39). The record from “Virginia” (Bousquet and Larochelle 1993: 114) is in error (see Hoffman et al. 2006: 19).
This species is known only from the type-locality cave in southeastern Kentucky (Barr 2004: 39). The record from “Virginia” (Bousquet and Larochelle 1993: 115) is in error (see Hoffman et al. 2006: 19).
This species is known only from the type-locality cave in northern Tennessee (Barr 2004: 39).
This species is known from several caves in the Rye Cove karst near Clinchport in southwestern Virginia (Barr 1981: 77).
This species is known only from two caves in Scott County, southwestern Virginia (Barr 2004: 39).
This species is known only from the type-locality cave, located two miles southwest of Springville, southern Indiana (Barr 2004: 27).
This subspecies is known from several caves in Monroe, Barren, Metcalfe, Adair, Cumberland, and Clay Counties in southern Kentucky and Jackson County in northern Tennessee (Barr 1985b: 124).
This subspecies is known from a number of caves in central Barren County, southern Kentucky (Barr 2004: 32).
According to Barr (2004: 32), intergrades between this subspecies and the nominotypical subspecies occur in Bowles Branch Cave in Barren County.
This subspecies is known from several caves in northeastern Metcalfe, northern Adair, and southern Green Counties, southern Kentucky (Barr 1985b: 126; Barr 2004: 32).
This subspecies is known from several caves in central Green and eastern Hart Counties, Kentucky (Barr 2004: 32).
This species is known only from the type-locality cave in central Kentucky (Barr 2004: 31).
This subspecies is found in several caves in Barren and Warren Counties, southern Kentucky (Barr 2004: 31).
According to Barr (2004: 31), intergrades between this subspecies and the nominotypical subspecies occur in caves in Warren County.
This subspecies is known from several caves in Hart, Edmonson, Barren, and Warren Counties in Kentucky (Barr 2004: 31).
This species is known from caves in Hardin and Hart Counties, central Kentucky (Barr 2004: 31).
This species is known only from the type-locality cave in central Kentucky (Barr 2004: 31).
This species is known from several caves in Hart, Edmonson, Metcalf, and Warren Counties, southern Kentucky (Barr 2004: 32).
The MCZ holds a specimen [# 7397], incorrectly labeled lectotype, of
This species is known from a few caves in Warren and Logan Counties, southern Kentucky (Barr 1985b: 123; Barr 2004: 31).
This species is known from a few caves in Smyth and Bland Counties, Virginia (Barr 2004: 19).
This species is known only from the type-locality cave in southwestern Virginia (Barr 2004: 19).
This species, the only one of the genus
This subspecies occurs in several caves in Montgomery, Cheatham, and Robertson Counties in northern Tennessee and Christian and Logan Counties in southern Kentucky (Barr 2004: 30).
This subspecies is known from a few caves in Logan and Simpson Counties, southern Kentucky, and Robertson County, northern Tennessee (Barr 2004: 30).
According to Barr (2004: 30), intergrades between this subspecies and the nominotypical subspecies occur in one cave in Robertson County, Tennessee.
This species is known from several caves in Montgomery County, northern Tennessee (Barr 2004: 31).
This species is known from several caves in Warren, Simpson, and Logan Counties, southern Kentucky, and Robertson and Sumner Counties, northern Tennessee (Barr 2004: 30).
This species is known from a few caves in Simpson and Warren Counties in southern Kentucky and Sumner County in northern Tennessee (Barr 2004: 30).
This subspecies is known two caves in Allen and Barren Counties, southern Kentucky (Barr 1985b: 128).
This subspecies is known from several caves from Hart and Metcalfe Counties westwards to eastern Warren County, Kentucky (Barr 2004: 30).
According to Barr (2004: 30), intergrades between the two subspecies of
This species is known only from a few caves in Greenbrier and Pocahontas Counties, eastern West Virginia (Jeannel 1949b: 75; Barr 2004: 20).
This species is known only from the type-locality cave in southeastern West Virginia (Barr 2004: 20).
This species is known only from two caves in Allegheny County, western Virginia (Barr 2004: 20).
This species is still known only from the type-locality cave in northern Virginia (Barr 2004: 20).
This species is known from a few caves in Giles County, western Virginia (Barr 2004: 20).
This species is found in several caves in Montgomery and Roanoke Counties, western Virginia (Barr 2004: 20).
This species is known only from the type-locality cave in east-central Kentucky (Barr 2004: 23).
This species is known from several caves in Estill, Powell, and Lee Counties, eastern Kentucky (Barr 2004: 23).
This species is known from several caves in Estill and Powell Counties, eastern Kentucky (Barr 2004: 23).
This species is known from several caves in Fentress, Overton, and Pickett Counties, northern Tennessee, and in McCreary and Wayne Counties, southern Kentucky (Barr 2004: 28).
This species occurs in several caves in DeKalb and Smith Counties, Tennessee (Barr 2004: 28).
This species is known from a several caves in DeKalb, Overtone, Putnam, Warren, White, Van Buren, and Grundy Counties, Tennessee (Barr 2004: 28).
This species is found in caves in Putnam and Overton Counties, northern Tennessee (Barr 2004: 28).
This species is known only from the type-locality cave in northern Tennessee (Barr 2004: 32).
This species is known from a few caves in Jackson County, northern Tennessee (Barr 2004: 32).
This species is known only from a few caves in Anderson County, eastern Tennessee (Barr 2004: 37).
This species is known only from the type-locality cave in eastern Tennessee (Barr 2004: 36).
This species is known from a few caves in Knox and Roane Counties, eastern Tennessee (Barr 1981: 55; Barr 2004: 36).
This species is known from two caves in Union County, northeastern Tennessee (Barr 2004: 37).
This species is known from several caves in Meade, Breckinridge, Hardin, Hart, and Larue Counties, Kentucky (Barr 2004: 26).
This species is known only from the type-locality cave (Barr 2004: 26).
This subspecies is known from several caves in Monroe, Lawrence, and Owen Counties, southern Indiana (Barr 2004: 26).
Barr (2004: 26) noted that intergrades between this subspecies and
This subspecies is known from a few caves in central Lawrence County, southern Indiana (Barr 2004: 26).
This species is known from several caves in Crawford, Washington, Orange, Lawrence, and Monroe Counties, southern Indiana (Barr 2004: 25).
This species is known from several caves in Crawford, Harrison, and Washington Counties, southern Indiana (Barr 2004: 25).
This species is known from several caves in Crawford, Orange, Lawrence, and Washington Counties, southern Indiana (Barr 2004: 26).
Two cave-inhabiting Appalachian species.
Valentine (1952) discussed the structural differences between the two species.
This species is known from a few caves in Pulaski County, southeastern Kentucky.
This species is known from two nearby caves in Putnam and Van Buren Counties, Tennessee.
One polymorphic cave-inhabiting species in the Appalachians.
Barr (1979a) revised the species and provided a key for the identification of its subspecies.
This subspecies is known from several caves in Breckinridge, Hardin, Hart, and Meade Counties in northwest and central Kentucky (Barr 1979a: 8).
This subspecies has been found in several caves in Allen, Simpson, Warren, and Logan Counties in southern Kentucky (Barr 1979a: 9).
This subspecies has been collected in several caves in Allen, Barren, Edmonson, Hart, and Warren Counties in central and southern Kentucky (Barr 1979a: 6).
Intergrades between this subspecies and the
This subspecies is known from several caves in Green, Hart, and Metcalfe Counties in central and southern Kentucky (Barr 1979a: 8).
Two Palaearctic species, one of them adventive in North America.
The species found in North America was covered in Lindroth’s (1961a: 194) monograph.
This Palaearctic subspecies is adventive in North America where it is known from Nova Scotia (NSMC) to Wisconsin (Messer 2010: 35), south to southern Pennsylvania (Bradford and Allegheny Counties, CMNH). The first inventoried specimen collected on this continent was found in the Montreal area in 1933 (Brown 1940a: 69).
The subspecies
Two troglobitic species restricted to southeastern Missouri.
Barr and Krekeler (1967) provided a description of the external structures and male genitalia of both species.
This species is known from two nearby caves, Friedman’s and Pleasant Valley Caves, in eastern Missouri.
This species is known from two nearby caves, Kohm’s and Sims Caves, in eastern Missouri.
One cave-inhabiting species in Kentucky.
Valentine (1952) lengthy described, neatly illustrated, and discussed the structural differences between members of
This species is known from several caves in Kentucky located as far north as Estill County and as far south as Fentress County [see Marsh 1969: Fig. 5].
One cave-inhabiting species in Kentucky.
Valentine (1952: 24-29) described, illustrated, and discussed the status of the species.
This subspecies is known from several caves in the Somerset area, southeastern Kentucky.
This subspecies is known only from the type-locality cave in southeastern Kentucky.
About 870 species arrayed in eight subgenera:
About 775 species in North America (23 species, of which three are adventive), mountains in Mexico (about five species), Oriental (two species in the Philippines), Palaearctic (about 725 species), and Afrotropical (about 25 species on Mont Elgon, Mont Meru, and Ethiopia) Regions. More than 95% of the species inhabit the Northern Hemisphere.
There is no published key for the identification of the species of this subgenus.
This species ranges from Newfoundland (Lindroth 1955a: 78-79, as
This species ranges from the Aleutian Islands in Alaska (Lindroth 1961a: 198) south to the Sierra Nevada in eastern California (Inyo and Tulare Counties, CAS; Casey 1918: 407, as
This species ranges from northwestern Washington (Hatch 1951: 113, as
This species ranges from Newfoundland and the coast of Labrador to northeastern Minnesota (Gandhi et al. 2005: 924), south to mountains in northern New York and New England [see Lindroth 1963a: Fig. 69]. The record from “Massachusetts” (Bousquet and Larochelle 1993: 120) is in error.
This species inhabits the North American Cordilleras ranging from British Columbia and southwestern Alberta (Lindroth 1963b: 201) south to western Montana (Russell 1968: 48) and the Sierra Nevada in California (Lindroth 1963b: 201). The record from “Colorado” (Bousquet and Larochelle 1993: 121) needs confirmation.
This species ranges from southern Yukon Territory (Lindroth 1961a: 200) south to mountains in northwestern Montana (Edwards 1975: 51), west-central Idaho (Boise County, CMNH), and western Oregon (Lane County, Foster F. Purrington pers. comm. 2009). The record from “Northwestern Territories” (Bousquet and Larochelle 1993: 121) needs confirmation.
This species is known only from Apache County (Donabauer 2010a: 41) in northeastern Arizona.
This species is known only from the type locality in the Bald Mountains between Greeneville, Tennessee and Asheville, North Carolina (Barr 1985b: 128).
This species is known only from the type locality, at the summit of Mount Mitchell where it is found in deep spruce and fir needle duff.
This species is found in caves in the Cumberland Plateau, from Rockcastle County in southeastern Kentucky southwest to Grundy County in southern Tennessee (Barr 1979b: 37).
This subspecies is known from Grandfather Mountain in Avery County and Three Top Mountain in Ashe County in western North Carolina (Barr 1979b: 43).
This subspecies intergrades with the
This subspecies ranges southwestwards from the Roan Mountain in Tennessee to the Black and Great Craggy Mountains at the edge of the Blue Ridge in western North Carolina (Barr 1979b: 44).
This subspecies is known from Grayson, Washington, and Lee Counties, southwestern Virginia, and Harlan and Letcher Counties, southeastern Kentucky (Barr 1979b: 36).
This subspecies is known from western Maryland, Virginia, and eastern West Virginia (Barr 1979b: 36). The record from Sassafras Mountain in northwestern South Carolina (Ciegler 2000: 44) is apparently in error (see Ciegler 2003: [1]) as well as the state record of “South Carolina” by Bousquet and Larochelle (1993: 121).
This species is found in the Black Mountains in Yancey, Buncombe, and McDowell Counties, North Carolina (Barr 1979b: 37).
This species is known from Carter County in northeastern Tennessee and from Mitchell County in western North Carolina (Barr 1979b: 37).
This subspecies is known from several specimens collected at the type locality in southwestern North Carolina.
This subspecies is restricted to the Pishah Ledge, which is the eastern arm of the Great Balsam Mountains, in western North Carolina (Barr 1979b: 45).
This subspecies is known from the Black and Great Craggy mountains and adjacent Blue Ridge, in Buncombe, McDowell, and Yancey Counties, western North Carolina (Barr 1979b: 46).
This species is known only from the San Jacinto Mountains in southeastern California.
This species is known only from the holotype.
This species is endemic to the Pinaleno Mountains in Graham County, southeastern Arizona (Donabauer 2010a: 39).
This species is known only from the holotype.
This species is known from northwestern California (Van Dyke 1945b: 101) and southern Oregon (Lane County, CMNH).
This species ranges along the Pacific Coast from southeastern Alaska (Lindroth 1961a: 196) to at least Marin County, central California (Kavanaugh and Erwin 1985: 177).
This species is known only from southwestern California (Jeannel 1927: 189; Donabauer 2010a: 39).
This European species is adventive in North America where it is known from the Queen Charlotte Islands (Kavanaugh 1992: 61) to north-central Idaho (LaBonte 1989: 17; Hatten et al. 2007: 359), south to northern Utah (Davis and Salt Lake Counties, CMNH) and west-central California [see Kavanaugh and Erwin 1985: Fig. 1]. The first inventoried specimen collected on this continent was found in North Creek, King County, Washington in 1925 (Kavanaugh and Erwin 1985: 171). The species is also adventive in Hawaii since 1998 (Liebherr and Takumi 2003).
This European species is adventive in North America where it is known from Nova Scotia (Majka et al. 2006: 603) to northern Wisconsin (Iron County, CMNH; Messer 2010: 35), as far north as the Abitibi region in Quebec (Paquin and Dupérré 2002: 87), south to northeastern West Virginia (Hampshire County, CMNH) and eastern Maryland (Queen Annes County, Foster F. Purrington pers. comm. 2009). The first inventoried specimen collected on this continent was found at Port Credit, southern Ontario, in 1965 (Bousquet et al. 1984: 215).
This European species is adventive in North America where it is known from Newfoundland (Lindroth 1955a: 82; Larson and Langor 1982: 593) to western Quebec (Larochelle 1975: 112), and from the New England area (Bousquet and Larochelle 1993: 121). The first inventoried specimen collected on this continent was found prior to 1863 (LeConte 1863b: 14) probably in Nova Scotia as recorded by Horn (1875: 131). The record from eastern Ontario (Hamilton 1889b: 94) is probably in error.
Forty-one species (46 species-group taxa) restricted to the Appalachian Mountains in North Carolina, Tennessee, South Carolina, and Georgia and currently placed in three species groups.
Barr (1979b) revised and provided a key to the species of this subgenus (22 species, 29 species-group taxa). Subsequently, 15 new species and two subspecies have been described by Barr (1985b), Dajoz (2005), and Donabauer (2005a, b).
This species is known only from the Plott Balsam Mountains, western North Carolina.
This species is known only from the type locality.
This species is known only from the type locality.
This species is known only from the type locality.
This subspecies is known only from the type locality.
This subspecies is known only from the type locality.
This subspecies is known from the central Great Smoky Mountains in North Carolina and Tennessee (Barr 1979b: 40).
This subspecies is endemic to the Nantahala Mountains in southwestern North Carolina (Donabauer 2005b: 88).
This species is known only from the type locality in the northern Nantahala Mountains, southwestern North Carolina.
This species is found in central and eastern Great Smoky Mountains in North Carolina and Tennessee (Barr 1979b: 40).
This species is found in central Great Smoky Mountains in North Carolina and Tennessee (Barr 1979b: 40).
This species is known only from the type locality.
This species is known only from the type locality.
This species is known from the Plott Balsam Mountains and Great Balsam Mountains in western North Carolina (Barr 1979b: 40).
This species is known only from the type locality.
This species is known only from the type locality.
This species is known only from the holotype collected in eastern Tennessee.
This subspecies is known only from Bull Cave in the Cades Cove Mountains, Great Smoky Mountains, in eastern Tennessee (Barr 1979b: 40).
This subspecies is known only from the original 28 specimens collected in a cave in the Appalachian Valley, 8 miles south of Kingston and ¼ mile west of the Tennessee River on the southeast side of a valley east of Huckleberry Ridge (Barr 1962a: 87).
This species is known only from the holotype.
This species is known only from the type locality in the Nantahala Mountains.
This species is known only from the original specimens collected in a large stream cavern at the north side of the Great Smoky Mountains.
This species is endemic to the Unicoi Mountains in southeastern Tennessee and western North Carolina (Barr 1979b: 40).
This taxon was first described as a subspecies but raised to species by Donabauer (2005b: 87).
This species is found in the central Great Smoky Mountains in Tennessee and North Carolina (Barr 1979b: 40).
This species is known only from the conifer forests in the eastern end of the Great Smoky Mountains in Haywood County, western North Carolina, and Cocke County, eastern Tennessee.
This species is known only from the type locality.
This species occurs in the Great Balsam Mountains in western North Carolina (Barr 1979b: 39).
This species is known only from the Cowee Mountains in southwestern North Carolina (Donabauer 2005a: 56).
This species is known only from Big Butt in the Nantahala Mountains, southwestern North Carolina (Donabauer 2005a: 57).
This species is known only from the type locality.
This species is known only from the type locality in the Plott Balsam Mountains, southwestern North Carolina.
This species is endemic to the western Pisgah Ridge (Donabauer 2005a: 57) in the Great Balsam Mountains.
This species has been found yet only in the Unicoi Mountains, between 4800-5000 feet, in North Carolina and Tennessee (Barr 1979b: 40).
This species is known only from Thunderhead Mountain at the junction of Swain County in North Carolina and Blount County in Tennessee.
This species is known only from the Toxaway Mountain in southwestern North Carolina.
This species is known only from the type locality in southwestern North Carolina.
This species is known from the central Great Smoky Mountains east to Plott Balsam Mountains in Sevier and Cocke Counties, eastern Tennessee, and in Haywood and Jackson Counties, western North Carolina (Barr 1979b: 38).
The range of this common species extends from the vicinity of Asheville, western North Carolina, to the mountains of northeast Georgia (Barr 1979b: 38) and northwestern South Carolina (Ciegler 2000: 44).
This species is found in the spruce-fir forests, between 3000-6500 feet, in the Great Smoky Mountains of North Carolina and Tennessee (Barr 1979b: 38; Donabauer 2009: 137).
This species is known only from the Unicoi Mountains, along the Tennessee-North Carolina border (Donabauer 2009: 138).
This species is known only from two localities in western North Carolina (Donabauer 2009: 136).
This species is known only from the type locality.
This species is known only from the type locality in southwestern North Carolina.
This species has been found at several locations in the Great Smoky Mountains and the Plott Balsams in western North Carolina (Barr 1979b: 38; Donabauer 2009: 131).
This species is known only from the type locality in the western Great Smoky Mountains on the border between Blount County, Tennessee, and Swain County, North Carolina.
This species is known at present from small mountain ranges in southwestern North Carolina (Barr 1979b: 37).
This subspecies is found in the Great Balsam and Cowee Mountains in western North Carolina (Barr 1979b: 37).
This subspecies is found in the Bald and Unaka Mountains of Tennessee and in the Black and Great Craggy Mountains of North Carolina (Barr 1979b: 37; Donabauer 2009: 138). Populations very similar to those of
Worldwide, with about 2,630 species arrayed in six subtribes:
Worldwide, with about 1,350 species. The number of genera admitted varies greatly depending on the authors. In this work, the species are arrayed in nine genera following Maddison (2012: 570):
About 50 species in North America (five species) and Asia (about 45 species) placed in two subgenera:
Toledano (2011) noted that
Six species in North America (five species) and the Altai Mountains in Russia (
Lindroth (1963b: 403-406) treated four of the five North American species, leaving
This species is restricted to the Queen Charlotte Islands and adjacent mainland (Kavanaugh 1992: 69).
This species ranges from the Kenai Peninsula in Alaska (Lindroth 1963b: 405) south to “California” (Hayward 1897: 131) and to the Sangre de Cristo Mountains in northeastern New Mexico (Ball 1966b: 30). The record from “Mexico” (Hayward 1897: 131) is probably in error.
The range of this species extends from the Alexander Archipelago (Mannerheim 1852: 298; Lindroth 1963b: 404) south at least to the Santa Cruz Mountains of the Coast Ranges (Casey 1918: 166, as
This species is known only from Utah.
This species was originally placed in the subgenus
This eastern species occurs from Newfoundland (Lindroth 1955a: 78, as
This species has long been confused with
Nine species in western North America.
Erwin and Kavanaugh (1981) revised the species and provided a key for their identification.
This species ranges from the southern part of the Alexander Archipelago, Alaska (Lindroth 1963b: 261) and west-central British Columbia south to central California along the coast mountain systems [see Erwin and Kavanaugh 1981: Fig. 23].
This species is known from several mountains in the Coast Ranges of Oregon (David R. Maddison pers. comm. 2012).
This species is known from four localities, the northernmost in southern British Columbia and the southernmost in the mid-Sierra Nevada [see Erwin and Kavanaugh 1981: Fig. 24].
This species ranges from southwestern Alberta and southeastern British Columbia south to the San Jacinto Mountains in southern California, including the Sierra Nevada and Cascade Range, and northern Colorado along the Rocky Mountains [see Erwin and Kavanaugh 1981: Fig. 25].
This species is known from a few localities along the Pacific Coast from southern Alaska near the Yukon Territory border to northern Washington [see Erwin and Kavanaugh 1981: Fig. 26].
This species corresponds to
This species is known from two localities, one on the eastern edge of the Central Plateau in eastern British Columbia, the other on an island on Puget Sound, Washington [see Erwin and Kavanaugh 1981: Fig. 27].
This species ranges from the lower Columbia River drainage in northwestern Idaho and southern Washington south to the Sierra Nevada and Coast Ranges in central California [see Erwin and Kavanaugh 1981: Fig. 20].
This species is endemic to the Sierra Nevada in California from Nevada County to Sequoia National Park [see Erwin and Kavanaugh 1981: Fig. 21].
This species ranges along the Cascade Range and Sierra Nevada from southern British Columbia to central California [see Erwin and Kavanaugh 1981: Fig. 22].
Northern Hemisphere, with 39 species in the arctic, subarctic, boreal, and temperate areas of the Nearctic (three species, one of them adventive) and Palaearctic (37 species) Regions.
Lindroth (1963b: 203-206) covered all three species found in North America, the adventive
This species is found from Alaska to northwestern Northwest Territories (Lindroth 1963b: 204). Fossil remnants of this species, believed to be 2.0-2.5 million years old, have been found in Greenland and Meighen Island (Bennike and Böcher 1990: 336; Böcher 1995: 23).
This European and North African subspecies is adventive in North America where it is known from Maine (Larochelle and Larivière 1990a: 28, 33, as
Two other subspecies, one from Morocco (
This species is known from Alaska, Yukon Territory, northeastern British Columbia, and west-central Alberta [see Morgan and Morgan 1979: Fig. 4; Morgan and Morgan 1981: map 4]. Fossil remnants of this species from the Pleistocene and early Holocene have been found in west-central Illinois, northeastern Wisconsin, Vermont, southern Ontario, and northwestern Ontario (see Ashworth and Schwert 1991: 512; Bajc et al. 1997: 691).
Worldwide, with about 1,250 species (Lorenz 2005: 215-236, as
Hayward (1897) reviewed the North American species but his work is now outdated. Lindroth (1963b) covered 191 (six of them in the key alone) of the 253 species found in North America (about 75% of the fauna).
Maddison (2012: 570) presented an entirely new classification of this genus based on molecular data analyses. He found support for monophyly of
Edwards (1975: 53) recorded a “
Northern Hemisphere, with 11 species in the Nearctic (nine species) and Palaearctic (two species:
There is no taxonomic revision of the North American species and such study is needed. Lindroth (1963b: 301-305) covered all but three (
This species is known only from the holotype collected in eastern California.
According to Erwin (1984a: 167), this species “is found in the Basin and Range Province of the western United States.”
This species ranges from Newfoundland (Lindroth 1955a: 56, as
As far as known, this species is confined to the west coast of southern Oregon (Hatch 1953: 88, as
The range of this eastern species extends from Cape Breton Island (Lindroth 1954c: 302) to eastern South Dakota (Kirk and Balsbaugh 1975: 17), south to east-central Louisiana (West Feliciana Parish, Igor M. Sokolov pers. comm. 2009), southwestern Mississippi (Drew A. Hildebrandt pers. comm. 2007), and northern Georgia (Fattig 1949: 17). The record from “Kansas” (Bousquet and Larochelle 1993: 134) needs confirmation.
The range of this western species extends from southern Yukon Territory (Lindroth 1963b: 304) south to the Sierra Nevada in California (Casey 1918: 50, as
This western species is found from south-central Alberta to Vancouver Island, north to the Skeena River drainage in central British Columbia (Lindroth 1963b: 305), south at least to central California in the Coast Ranges (Casey 1918: 53, as
This species ranges from Newfoundland (Lindroth 1955a: 56) to the foothills of the Rockies in Alberta and northeastern British Columbia (Lindroth 1963b: 302), south to southeastern Wyoming (Hayward 1897: 60, as
This species has passed under the name
This species is known only from the type locality in the Sierra Nevada in California.
Five North American species, two in the east and three in the west.
Lindroth’s (1963b: 212-229) key to
Based on molecular data analyses, Maddison (2012: 568) concluded that this subgenus is closely related to
This species is known from southwestern Oregon (Hatch 1953: 88) to southern California “in the higher mountain cañons or valleys between the ranges” (Fall 1901a: 42).
This species is known from southwestern Oregon (Josephine County, James R. LaBonte pers. comm. 1992) to the southern part of California where it is “widely distributed” (Fall 1901a: 42). The record from northwestern Montana (Edwards 1975: 51) needs confirmation.
This species is known only from Del Norte (Maddison 2012: Supplementary content Table S1), Marin (Casey 1924: 29, as
This species ranges from Newfoundland (Lindroth 1955a: 50) to the Laurentides region in Quebec (Larochelle 1975: 60), south to southwestern (Hardin County, CMNH) and eastern Tennessee (Johnson, Morgan, Roane, and Union Counties, CMNH).
The range of this eastern species extends from Nova Scotia (Lindroth 1954c: 302) to west-central Indiana (Blatchley 1910: 79), including southeastern Michigan (Wayne County, CMNH), south to northwestern Arkansas (Newton County, CNC, UASM), northeastern Mississippi (Tishomingo County, Drew A. Hildebrandt pers. comm. 2010), and southern Georgia (Torres and Ruberson 2006: 31).
Twenty-three species in the Nearctic (ten species), Neotropical (one species also present in southern Arizona), Palaearctic (12 species), and Oriental (one species,
Lindroth (1963b: 241-246, as
This species is known from eastern South Dakota to southwestern Nebraska, east to eastern Iowa [see Maddison and Arnold 2009: Fig. 9]. Two specimens simply labeled “Missouri” and “Wisconsin” (Maddison and Arnold 2009: 60) are known.
This species is known from a few localities in southern British Columbia (Lindroth 1963b: 242), northwestern Washington (Maddison 2012: Supplementary content Table S1), “Idaho” (Maddison 1993: 161), western Montana (Russell 1968: 49), and southwestern Wyoming (Hayward 1897: 51).
This species is known from a few localities in southeastern California and southern Arizona (Lindroth 1963b: 245; Dajoz 2007: 19).
This species is found east of the Rocky Mountains from New Brunswick (Queens County, Reginald P. Webster pers. comm. 2008) to southeastern Alberta (Lindroth 1963b: 245), south to southeastern Colorado (Maddison 1985: 111), northwestern (Jones County, CNC) and east-central (Riley 2011) Texas, and central Florida (Peck and Thomas 1998: 18).
The range of this species extends from southwestern and north-central Pennsylvania (Westmoreland and Lycoming Counties, Robert L. Davidson pers. comm. 2008) to Alberta (Lindroth 1963b: 244), south to central New Mexico (Fall and Cockerell 1907: 157; Socorro County, UASM), southern Texas (Snow 1906a: 141; Casey 1918: 12, as
This species is known from southern Arizona (Lindroth 1963b: 246; Dajoz 2007: 21) and northern Mexico (Bates 1891a: 263).
This species is known from the Gila River drainage in southern Arizona.
This taxon has been listed in synonymy with
This species ranges from southern New Hampshire south to northern Florida [see Maddison and Arnold 2009: Fig. 9], west to eastern Texas (Hardin County, Robert L. Davidson pers. comm. 2012), including eastern Louisiana (East Feliciana Parish, Igor M. Sokolov pers. comm. 2009).
This species is known from a few scattered localities from Connecticut (Lindroth 1963b: 243; Krinsky and Oliver 2001: 82) and northern Vermont (Lamoille County, Ross T. Bell pers. comm. 2008) to western North Dakota (McKenzie County, Donald P. Schwert pers. comm. 1989), south at least to “Kansas” (Hayward 1897: 50) and “Kentucky” (Lindroth 1963b: 243).
This species is known from eastern Washington (Hatch 1953: 80) to central California (Lindroth 1963b: 245).
Northern Hemisphere, with 17 species in the Nearctic (11 species) and Palaearctic (nine species) Regions. Three species are Holarctic.
Maddison (1993) revised the species of this subgenus and provided a key for their identification.
This Holarctic species is known from the region of Lake Baikal in Siberia and Alaska, from the Brooks Range south to the Alaska Peninsula [see Maddison 1993: Fig. 264]. Fossil remnants from a Plio-Pleistocene sequence have been found in northwestern Greenland (Böcher 1995: 24).
This species is known only from west-central Saskatchewan and eastern Alberta [see Maddison 1993: Fig. 261]. The record from Churchill, Manitoba (Elias 1984: 142) is based on misidentified
A common species ranging from Newfoundland to southwestern British Columbia, north to central Yukon Territory and the Ungava Bay in Quebec, south to northern Iowa and southwestern Pennsylvania [see Maddison 1993: Fig. 265]. The records from “Georgia,” “Arkansas,” “Oregon” and Vancouver Island (Hayward 1897: 46) are probably in error; that from “Kentucky” (Maddison 1993: Fig. 265) needs confirmation. Lindroth’s (1963b: 237) record from Washington is based on a misidentified
This Holarctic species ranges in the Palaearctic Region from northeastern Europe to Kamchatka, and in the Nearctic Region, from Alaska east to Churchill, Manitoba, and south to central Saskatchewan [see Maddison 1993: Fig. 262a].
This western species ranges from southeastern British Columbia to Vancouver Island, south to east-central California [see Maddison 1993: Fig. 269].
This species ranges from Cape Breton Island to western Alaska, south to northwestern California, south-central Colorado, northern Kansas, southeastern Louisiana (East Baton Rouge Parish, Igor M. Sokolov pers. comm. 2009), northern Alabama, and northern Georgia [see Maddison 1993: Fig. 273]. The records from “Florida,” “Arkansas,” and “Texas” (Hayward 1897: 44) are probably in error.
This Holarctic species ranges in the Palaearctic Region from northernmost Scandinavia to Kamchatka, south to Mongolian Peoples’ Republic. In North America, it occurs commonly in the northwest from Alaska to the Anderson River; it is also found at scattered localities in Washington, northern Oregon, Idaho, and Wyoming, and in western Canada east to central Saskatchewan [see Maddison 1993: Fig. 267a].
This transamerican subspecies ranges from western Newfoundland (Lindroth 1955a: 47) and Labrador to western Alaska, south to west-central British Columbia, southeastern New Mexico, southern Wisconsin, southwestern Michigan, central New York, and southeastern New Brunswick [see Maddison 1993: Fig. 272].
The range of this subspecies extends from central Washington to southwestern Alberta, south to northern New Mexico [see Maddison 1993: Fig. 272]. One specimen labeled “Cal.” is known (Maddison 1993: 191). Fossil remnants from a Plio-Pleistocene sequence have been unearthed in northwestern Greenland and Banks Island (Böcher 1995: 23).
This species has long been known in the North American literature under the name
This western species is found from southwestern British Columbia, including Vancouver Island, to western Idaho, south to southwestern California [see Maddison 1993: Fig. 270]. The records from Wyoming (LeConte 1878a: 465), Colorado (Wickham 1902: 232; Lindroth 1963b: 236), and “Montana” (Hayward 1897: 46) are probably in error.
This species shows a disjunct distribution. In the east, it ranges from Nova Scotia (Majka et al. 2007: 8), Maine, and southern Quebec south to the District of Columbia and Kentucky; in the west it occurs from western North Dakota and Saskatchewan to southwestern British Columbia, north to northern Northwest Territories and southwestern Alaska (Elias 1988: 41) [see Maddison 1993: Fig. 268]. I have also seen one specimen from northwestern Colorado (Eagle County, CMNH). The records from “Missouri,” “Arkansas” (Hayward 1897: 47), southwestern Iowa (Wickham 1911b: 6), “Washington” and “Oregon” (Hatch 1953: 81) need confirmation; that from “California” (Hayward 1897: 47) is probably in error.
This species ranges from the Queen Charlotte Islands (Kavanaugh 1992: 62) south to San Francisco, California; it is mainly found on the Pacific Coast though a few inland localities are also known [see Maddison 1993: Fig. 271]. The record from Mono County in eastern California (Dajoz 2007: 17) needs confirmation.
Three North American species.
Lindroth (1963b: 249-250) recognized two species,
This species ranges from southern New Brunswick (Webster and Bousquet 2008: 16) to southwestern Wisconsin (Richland County, Peter W. Messer pers. comm. 2008), south to eastern Texas (Sabine County, Brian Raber pers. comm. 2010; Riley 2011) and central Florida (Peck and Thomas 1998: 18). The record from “Minnesota” (Bousquet and Larochelle 1993: 126) needs confirmation.
This species ranges from southern British Columbia, including Vancouver Island (Lindroth 1963b: 250), south to southern California (LeConte 1852a: 186) and southwestern Utah (Casey 1918: 14, as
If the lectotypes currently associated with the labels for
The range of this species extends from Nova Scotia (Majka et al. 2007: 7) to the Rocky Mountains in Alberta (Lindroth 1963b: 250, as
Nine North American species, one of them extending into northern Mexico.
Maddison (2008) revised the species and provided a key for their identification. Some of the species are very difficult to recognize on external characters and the male genitalia must be examined.
This species ranges from Nova Scotia to northwestern Wisconsin, south to northern Arkansas, central Mississippi (Madison County, Drew A. Hildebrandt pers. comm. 2008), northern Alabama, and central North Carolina [see Maddison 2008: Fig. 21].
This species ranges from New England and northeastern New York south to northeastern Kentucky [see Maddison 2008: Fig. 20A].
The range of this species extends from Newfoundland to southwestern Northwest Territories, south to northeastern California, northern Colorado, north-central Arkansas, and South Carolina [see Maddison 2008: Fig. 18]. The record from “Mississippi” (Bousquet and Larochelle 1993: 126) probably refers to another species of the subgenus.
This species ranges from Newfoundland to southeastern Quebec, south to central New Hampshire [see Maddison 2008: Fig. 22]
This species ranges from Cape Breton Island to east-central Missouri, south to southwestern Mississippi and southern North Carolina along the Appalachian Mountains [see Maddison 2008: Fig. 19].
This species ranges from the Roanoke River drainage in Virginia south to southeastern Mississippi in the Piedmont, east to eastern North Carolina [see Maddison 2008: Fig. 23]
This species ranges from Cape Breton Island to eastern Illinois, south to North Carolina along the Appalachian Mountains [see Maddison 2008: Fig. 23]. One old specimen simply labeled from Wisconsin is known (Messer 2010: 35). The records from “Iowa,” “Kansas” (Hayward 1897: 56), and “Michigan” (Bousquet and Larochelle 1993: 126) need confirmation; that from Churchill in northern Manitoba (Elias 1984: 142) is in error (Maddison 2008: 181); those from “Wyoming,” “Texas” (Hayward 1897: 56), “Arkansas,” and “Mississippi” (Bousquet and Larochelle 1993: 126) are probably in error.
This species ranges from west-central Wisconsin to south-central Montana, south to north-central New Mexico, southern Texas, northwestern Tamaulipas in Mexico, eastern Louisiana (East Feliciana Parish, Igor M. Sokolov pers. comm. 2009), and southern Mississippi, east to west-central Indiana [see Maddison 2008: Fig. 24].
This species ranges from New Brunswick (Webster and DeMerchant 2012: 5) and southern Quebec (André Larochelle and Reginald P. Webster pers. comm. 1997, 2009) south to northern Georgia (Choate and Choate 1995: 371) and northwestern South Carolina (Ciegler 2003: [1]) along the Appalachian Mountains, east to northeastern Massachusetts [see Maddison 2008: Fig. 20B).
Two species, one along the Pacific Coast of North America, the other (
Kavanaugh and Erwin (1992: 317) emended Lindroth’s (1963b) key to
This taxon is ranked as a distinct genus by many authors, with
This species is known only from two localities on the Palos Verdes Peninsula along the coast of southern California (Kavanaugh and Erwin 1992: 315).
Northern Hemisphere, with five species in the Nearctic (one species) and Palaearctic (four European and northern African species, one of them endemic to the Canary Islands) Regions.
Lindroth (1963b: 265) treated the North American species.
This species is known from the Whitefox River drainage in east-central Saskatchewan and several localities in Alberta, as far north as the Fort Vermilion area (Bousquet 1987a: 120; CNC).
About 40 species in the Nearctic (one Holarctic species), Palaearctic (about 30 species), and Afrotropical (nine species) Regions.
Lindroth (1963b: 342-343) covered the species found in North America.
This Holarctic species is found in Kazakhstan, Siberia, Mongolia, Liaoning in China, the Korean Peninsula, and Japan (Marggi et al. 2003: 255), and from mainland Alaska (Lindroth 1963b: 343) to Newfoundland (Lindroth 1955a: 58-59), south to Connecticut (Litchfield County, William L. Krinsky pers. comm. 2008; Lindroth 1963b: 343), north-central Ohio (Lee 1994: 59), southeastern South Dakota (Kirk and Balsbaugh 1975: 18), and northern New Mexico (Casey 1918: 71, as
About 65 species in the Nearctic (21 species, three of them adventive), Neotropical (four species in Middle America shared with North America), Palaearctic (about 50 species), and Afrotropical (one species in Ethiopia,
Lindroth (1963b: 330-342, as
This Holarctic species ranges from Scandinavia to eastern Siberia and Mongolia (Marggi et al. 2003: 261) in the Palaearctic Region and from the Chukchi coast in Alaska to the Hudson Bay coast in northern Manitoba (Lindroth 1963b: 325); it also occurs quite isolated in the southern parts of the Rocky Mountains in Colorado and New Mexico (Lindroth 1969a: 1114). Fossil remnants of this species, dated between about 16,700 and 18,100 years B.P., have been unearthed in southeastern Iowa (Baker et al. 1986: 96); others, older than 33,000 years B.P., in southwestern Ontario (Warner et al. 1988: 35).
This species inhabits the North American Cordilleras from southwestern Alberta and southern British Columbia (Lindroth 1963b: 318, as
This species is known from southeastern Wyoming (Lavigne 1977: 44), south-central Colorado (Wickham 1902: 233; Lindroth 1963b: 316), and “New Mexico” (Casey 1918: 74) west to Guadaloupe Island off the Pacific Coast of Baja California (Hayward 1897: 76). The record from “Montana” (Bousquet and Larochelle 1993: 135) needs confirmation; that from “Oregon” (Notman 1919b: 227) is in error according to Lindroth (1963b: 316).
This species is widely distributed in the North American Cordilleras from southern British Columbia south to central California along the Sierra Nevada (Lindroth 1963b: 317), southeastern Arizona (Cochise County, UASM, Ken Karns pers. comm. 2009), and “New Mexico” (Casey 1918: 65, as
This species, the only one of the group occurring in eastern North America, is known from southernmost Ontario (Lindroth 1963b: 319) to “New Jersey,” south to south-central North Carolina (Hoffman 1982: 146), northeastern Georgia (Horn and Ulyshen 2009: 121), eastern Alabama (Tallapoosa County, UASM, Drew A. Hildebrandt pers. comm. 2009), and east-central Mississippi (Lowndes County, Drew A. Hildebrandt pers. comm. 2008).
This species is known from a few localities in the Sierra Nevada north of Yosemite National Park (Lindroth 1963b: 318; Maddison 1985: 113).
This species ranges from Vancouver Island (Lindroth 1963b: 315) south to southern California (LeConte 1852a: 190) where it is “widely distributed” (Fall 1901a: 42). The records from Colorado (Wickham 1902: 233), New Mexico (Fall and Cockerell 1907: 157), central Arizona (Griffith 1900: 565), Utah (Knowlton 1939: 2), northern Idaho (Hatch 1953: 90), and northeastern Kansas (Popenoe 1878: 79) are probably in error.
This European species is adventive in North America where it is known from Newfoundland (Lindroth 1955a: 61, as
This European subspecies is adventive in North America where it is known from New Brunswick and Nova Scotia (Bousquet 1992a: 504; Majka 2005: 536) and from one specimen discovered in 2003 in Grant County, Washington (Robert L. Davidson pers. comm. 2008). The first inventoried specimen collected on this continent was found near Halifax, Nova Scotia in 1967 (Bousquet 1992a: 504).
This Holarctic subspecies ranges from Scandinavia to northeastern Siberia (Motschulsky 1845a: 27) and Mongolia (Marggi et al. 2003: 262) and in North America from Alaska (Lindroth 1963b: 339) to Newfoundland (Bousquet 1987a: 120), south to Virginia (Hoffman et al. 2006: 20), east-central Iowa (Iowa County, MCZ), southern New Mexico (Fall and Cockerell 1907: 157), southwestern Nevada (Esmeralda County, CMNH), and “California” (Cooper 1976: 163).
Five more subspecies are recognized within the Palaearctic fauna.
This subspecies is known only from numerous specimens collected at the type locality.
This Holarctic subspecies is known from Scandinavia to eastern Siberia (Marggiet al. 2003: 262) and from Alaska (Lindroth 1963b: 334) to Newfoundland (Lindroth 1955a: 60-61), south to “Rhode Island” (Sikes 2003: 7), southwestern Pennsylvania (Allegheny County, CMNH), “Nebraska” (Hayward 1897: 81, as
This subspecies is known only from a few localities on mainland Alaska.
Besides the nominotypical subspecies found in eastern Siberia, two more subspecies have been described from Japan and Szechwan, China.
The range of this species extends from the Ontario Peninsula (Bousquet 1987a: 120) to southern British Columbia, north to western Alaska (Lindroth 1963b: 337-338), south to northern Oregon (Gilliam and Morrow Counties, CMNH; Hatch 1953: 90; Lindroth 1963b: 337), southern Arizona (Dajoz 2007: 21), western Texas (Jeff Davis County, MCZ), and northwestern Pennsylvania (Warren and Forest Counties, CMNH). The species has been reported also from “Mexico” (Casey 1924: 35, as
The range of this subspecies is disjunct. It occurs in the east along the coast of Newfoundland, Saint Pierre and Miquelon (Lindroth 1955a: 63), the Maritimes, Magdalen Islands (Lindroth 1963b: 340), and the Saint Lawrence Estuary (Larochelle 1975: Fig. 169) and in the west from the Mackenzie River in central Northwest Territories south to northern New Mexico (Taos County, UASM), including southwestern North Dakota (Tinerella 2003: 635) [see Lindroth 1963a: Fig. 66].
This subspecies is confined to the west coast ranging from the Queen Charlotte Islands (Kavanaugh 1992: 66) south at least to northern California (Fall 1922c: 171; Hatch 1953: 91) [see Lindroth 1963a: Fig. 66].
This Palaearctic subspecies is adventive in North America where it is known from Newfoundland (Lindroth 1955a: 62, as
This species has been known for a long time under the name
This species ranges from “California” to “Texas,” south at least to Oaxaca, Mexico (Maddison and Swanson 2010: 29).
This species ranges from South Dakota (Maddison and Swanson 2010: 28) and eastern Wyoming (Lavigne 1977: 44, as
This species is known from southern Colorado and northern New Mexico (Maddison and Swanson 2010: 29).
This species ranges from “Oregon” (Maddison and Swanson 2010: 27) to western Nebraska (Keith County, Robert L. Davidson pers. comm. 2012), south to western Texas (Jeff Davis and Presidio Counties, CNC) and southern California (Maddison and Swanson 2010: 27).
This species ranges from “Wyoming” south to northern New Mexico and northeastern Arizona (Maddison and Swanson 2010: 28).
This species ranges from Newfoundland (Lindroth 1955a: 59) to Alaska (Maddison and Swanson 2010: 27), south to southern California (LeConte 1852a: 190; Maddison 1985: 114), “Utah,” “Colorado” (Maddison and Swanson 2010: 27), northern Michigan (Hubbard and Schwarz 1878: 629; Dunn 1985a: 10; CNC), and central New York (Notman 1928: 215). The record from South Dakota (Kirk and Balsbaugh 1975: 17) refers to
Maddison and Swanson (2010: 27) stated that specimens of this species from western British Columbia, western Oregon, and California represent another form, with no consistent differences in COI or 28S from the typical form but with clearly different mentum shape. They added that
About 15 species in the Nearctic (one northwestern species), Oriental (one species,
Lindroth (1963b: 307-310) covered the species found in North America.
This subspecies is known only from the Rocky Mountains and foothills in British Columbia and Alberta, as far north as northern British Columbia near the Yukon Territory border (Lindroth 1963b: 309). The record from southwestern Alaska (Elias 1988: 41) possibly refers to the nominotypical subspecies and needs confirmation.
This subspecies is known only from southwestern Yukon Territory and Alaska from the Gulf Coast to beyond the arctic circle (Lindroth 1963b: 308). Fossil remnants of this species, dated between about 10,000 and 11,000 years old, have been found in southern Quebec (Mott et al. 1981: 146) and southern Ontario (Pilny et al. 1987: 620).
Another subspecies,
Northern Hemisphere, with about 30 species in the Nearctic (six species) and Palaearctic (27 species) Regions. Two species (
Lindroth (1963b: 326-330, as
This species ranges from north-central Ontario (CNC) to the Arctic Plains of Alaska (Lindroth 1963b: 326), south to San Bernardino County in southeastern California (Dajoz 2007: 20), northeastern Arizona (Apache County, UASM; Wickham 1896a: 157), and southern New Mexico (Fall and Cockerell 1907: 157). The record from “Kansas” (Hamilton 1894a: 7) is probably in error.
This Holarctic species is found from eastern Siberia (Marggi et al. 2003: 243) to the Anderson River delta in northern Northwest Territories (Lindroth 1963b: 311).
This eastern species is found from “New Brunswick” (Larochelle and Larivière 1990a: 28) to northern Minnesota (Gandhi et al. 2005: 925), south to northeastern Iowa (Purrington and Larsen 1997: 50), southwestern Ohio (Dury 1910: 66), and southwestern Pennsylvania (Hayward 1897: 80; Lindroth and Freitag 1969: 336). The record from northeastern Georgia (Fattig 1949: 17) is probably in error.
This species is known only from the type locality in western Nevada.
The range of this species extends from the Ungava Bay area (Larochelle 1975: Fig. 173) in northern Quebec to the Arctic Plains of Alaska (Lindroth 1963b: 329), south to east-central Utah (Grand County, UASM), southwestern Colorado (Elias 1987: 632), southern South Dakota (Kirk and Balsbaugh 1975: 18), and the Saint Lawrence River drainage in southern Quebec (Larochelle 1975: Fig. 173). The record from “Michigan” (Bousquet and Larochelle 1993: 137) needs confirmation. Fossil remnants, dated from about 10,400 and 21,500 years B.P., have been unearthed in eastern Minnesota, northeastern Wisconsin, Iowa, northern Illinois (Schwert 1992: 77), northeastern Pennsylvania (Barnosky et al. 1988: 178), and Cape Breton Island in Nova Scotia (Miller 1997: 250).
This Holarctic species is found from the northern regions of European Russia to eastern Siberia (Marggi et al. 2003: 244) and in North America from the Arctic Plains in Alaska (Lindroth 1963b: 330) to Contwoyto Lake in western Nunavut (Shpeley and Pilny 1995: 233).
Northern Hemisphere, with about 35 species in the Nearctic (seven species, one of them adventive) and Palaearctic (about 30 species) Regions. Two species (
Lindroth (1963b) covered all but one (
This Holarctic species ranges in North America from the Seward Peninsula in Alaska (Lindroth 1963b: 320) to Newfoundland (Lindroth 1955a: 57) and Greenland (Böcher 1988: 12), including the Aleutian and Kodiak Islands, south to mountains in New England (Lindroth 1963b: 320), southeastern upper peninsula of Michigan (Mackinac County, UMAA), northern Wisconsin (Bayfield County, MCZ), northeastern Minnesota (Kamal J.K. Gandhi pers. comm. 2008), southern New Mexico (Fall and Cockerell 1907: 157), and southeastern Arizona (Dajoz 2007: 21; Lindroth 1969a: 1114). The records from “Massachusetts” and “Pennsylvania” (Bousquet and Larochelle 1993: 136) are in error.
This species is listed as
This eastern species is found from New Brunswick (Webster and Bousquet 2008: 17) to “North Dakota” (Donald P. Schwert pers. comm. 1989), south to Big Bend National Park in western Texas (Dajoz 2007: 23) and Tennessee (Cannon, Clay, Fayette, Macon, Madison, Overton, Smith, Wayne, and Wilson Counties, CMNH), including northeastern New Mexico (Fall and Cockerell 1907: 157, as
This species has been known for a long time under the name
The range of this western species extends from Vancouver Island (Lindroth 1963b: 323) to western Montana (Russell 1968: 53; Edwards 1975: 53), north to central British Columbia (Lindroth 1963b: 323), south to southern California (Hayward 1897: 79; Fall 1901a: 42; Moore 1937: 6). The record from “New Mexico” (Bousquet and Larochelle 1993: 136) is probably in error.
This Palaearctic species is adventive in North America where it is known from Newfoundland (Lindroth 1955a: 58) to northeastern Wisconsin (Forest County, CMNH; Messer 2010: 36), south to northern Ohio (Lee 1994: 58) and Massachusetts (Lindroth 1955a: 58). The records from “Rhode Island” and “Virginia” (Bousquet and Larochelle 1993: 136) need confirmation. The first inventoried specimen collected on this continent was found in Ottawa, Ontario in 1891 (Lindroth 1963b: 323).
This species is known from western Texas (Jeff Davis County, Ken Karns pers. comm. 2009), southern New Mexico (Fall and Cockerell 1907: 157), southern Arizona (Casey 1918: 67), southeastern California (Dajoz 2007: 20), and “Mexico” (Casey 1918: 67).
This species ranges from northwestern Ohio (Holeski and Graves 1982: 216) to southern Manitoba (Lindroth 1963b: 326), south to southeastern Arizona (Maddison 1985: 114; Dajoz 2007: 21), northwestern New Mexico (Casey 1918: 67 as
This Holarctic species is found from Scandinavia to eastern Siberia (Marggi et al. 2003: 272) and in North America from the Arctic Plains in northern Alaska (Lindroth 1963b: 321) to the north shore of the Saint Lawrence Estuary (Larochelle 1975: Fig. 183), south to east-central British Columbia (Lindroth 1963b: 321).
This species is listed as
Northern Hemisphere, with 24 species in the Nearctic (6 species) and Palaearctic (18 species) Regions arrayed in three species groups, two of these groups being exclusively Eurasian. One species from India,
Lindroth (1963b: 312-314, as
This species is found in the North American Cordilleras from southeastern Alberta and southern British Columbia south to the Sierra Nevada in California (Lindroth 1963b: 313) and to central Colorado (Wickham 1902: 233, as
This species has been known until the 1960s under the name
This species is known so far only from the type series collected in northeastern California.
This western species is known from southwestern Alberta and southern British Columbia (Lindroth 1963b: 312) south at least to northeastern California (Modoc County, CNC; Hayward 1897: 69), Nevada (White Pine County, CMNH), northern Utah (Rich County, UASM), and northern New Mexico (Taos County, UASM; Hayward 1897: 69; Maddison 2012: Supplementary content Table S1). The record from “Nebraska” (Hayward 1897: 69) probably originated from the original statement about the type locality.
This species is found along the North American Cordilleras from northern Washington (Hatch 1950: 101, as
According to Erwin (1984a: 178), this species occurs in the foothills of the Sierra Nevada of California.
This species is known only from the original six specimens collected in “Nevada.”
One North American species along the Pacific Coast.
The species is included in Lindroth’s (1963b: 224) key to the Canadian
This species ranges along the Pacific Coast from the southern tip of Vancouver Island (Bousquet 1987a: 121) to southern California (LeConte 1879d: 509; Fall 1901a: 9).
Northern Hemisphere, with 11 species in the Nearctic (six species) and Palaearctic (six species) Regions. One species (
Lindroth (1963b: 382-387) covered four of the North American species in his monograph. A revision of the Nearctic species is needed.
This species is known from the Sierra Nevada (Casey 1918: 153, as
The range of this species extends from Newfoundland (Lindroth 1955a: 73) to east-central Alaska (Lindroth 1963b: 387), south to Nechako River in central British Columbia, northern New Mexico (Fall and Cockerell 1907: 157), west-central Minnesota (Gandhi et al. 2005: 924), the Adirondack Mountains in northeastern New York (Notman 1928: 218), and New England (Lindroth 1963b: 387). The record from “Washington” (Bousquet and Larochelle 1993: 146) is probably in error.
As far as known, this species has been recorded from the type locality and Frenchglen, Harney County, in Oregon (Hatch 1953: 98).
This species ranges from southeastern Pennsylvania (Delaware and Lebanon Counties, Robert L. Davidson pers. comm. 2008) to southeastern Minnesota (Gandhi et al. 2005: 925), south to central Texas (Bosque County, CMNH) and eastern Tennessee (Sevier County, CNC). The records from “New Jersey” and “North Carolina” (Bousquet and Larochelle 1993: 146) need confirmation.
This species is found from southern Manitoba (Lindroth 1963b: 386) to southeastern Washington (Hatch 1953: 98), including southern Saskatchewan and Alberta (Lindroth 1963b: 386), south at least to Santa Barbara County in southwestern California (Maddison 1985: 113), southern Arizona (Dajoz 2007: 21), and southern Colorado (LeConte 1879d: 509; Wickham 1902: 234).
This subspecies ranges from the southwestern edge of Lac Winnipeg in southern Manitoba to Vancouver Island (Hayward 1897: 119), north to the Brook Range in Alaska (Lindroth 1963b: 385), south to the Baja California Peninsula (Horn 1894: 308) and southern New Mexico (Fall and Cockerell 1907: 157). The records from Kansas (Snow 1903: 193; Knaus 1905a: 218) need confirmation.
Three other subspecies, including the nominotypical one, are known from the Palaearctic Region (Toledano 1999: 205-210).
This subspecies ranges from Newfoundland (Lindroth 1955a: 72) to the Rocky Mountains in Alberta (Lindroth 1963b: 384), south to central Colorado (Wickham 1902: 235), “Texas” (Lindroth 1963b: 384), northern Georgia (Fattig 1949: 18), and east-central South Carolina (Ciegler 2000: 48).
Six Neotropical species of which three extend into southwestern North America.
Perrault (1982a) revised the species except
This species is known from the Baja California Peninsula and “Arizona” (Hayward 1897: 121).
This species ranges from southeastern Arizona and southwestern New Mexico (Hidalgo County, CMNH) south to the Isthmus of Tehuantepec in southern Mexico [see Perrault 1982a: Fig. 64].
This species is found from Arizona and New Mexico south to Oaxaca in Mexico [see Perrault 1982a: Fig. 64].
Western Hemisphere, with nine species in North America (nine species) and northern Neotropical (three species in Cuba and Mexico, all shared with North America) Regions.
Lindroth (1963b: 376-381, as
This species ranges from southwestern Quebec (CNC) to “North Dakota” (Donald P. Schwert pers. comm. 1989), south to east-central Texas (Casey 1918: 128; Riley 2011) and the Florida Panhandle (Peck and Thomas 1998: 18); also recorded from Cuba (Jacquelin du Val 1857: 23). The records from Arizona (Wickham 1896a: 157; Hayward 1897: 122) and “Colorado” (Wickham 1902: 235) need confirmation.
This species is known from central Wyoming (Natrona County, CMNH) south to northwestern New Mexico (Casey 1918: 132) and southern Arizona (Casey 1918: 133, as
This species is known from eastern Washington (Hatch 1953: 97) to central Wyoming (Natrona County, CMNH), south at least to the Sierra Nevada in east-central California (Blaisdell 1902: 77).
This species occurs from southwestern New Brunswick (Larochelle and Larivière 1990a: 29, 34) to southern British Columbia (Lindroth 1963b: 381), south to southeastern California (Casey 1918: 130, as
The range of this species extends from southern Newfoundland (Lindroth 1955a: 72) to southern Saskatchewan (Ronald R. Hooper pers. comm. 1990), south to northern Nebraska (Cherry County, Foster F. Purrington pers. comm. 2010), “Illinois” (Lindroth 1955a: 72), and northeastern West Virginia (Tucker County, CMNH). The record from the Atlantic Coast in South Carolina (Kirk 1969: 10; Ciegler 2000: 48) needs confirmation.
This species has been collected in western Texas (Brewster County, CMNH), southern Arizona (Cochise, Pima, and Pinal Counties, CMNH; Casey, 1924: 42), and in Baja California Sur (CMNH).
This species is found from western Ontario to Vancouver Island (Lindroth 1963b: 379), north to northern Yukon Territory (Maddison 1985: 114) and southern Northwest Territories (Lindroth 1963b: 379), south to southern California (Fall 1901a: 42; Andrews et al. 1979: 28), southern Arizona (Wickham 1898: 300; Griffith 1900: 565; Snow 1907: 142), central New Mexico (Ellis et al. 2001: 16), southern South Dakota (Kirk and Balsbaugh 1975: 19), and northwestern Minnesota (Polk County, CMNH; Kittson and Roseau Counties, Peter W. Messer pers. comm. 2009). The record from “Michigan” (Bousquet and Larochelle 1993: 145) needs confirmation.
This species is known only from Lake County in the California Coast Ranges.
This species is found from Newfoundland (Lindroth 1955a: 70-71) to central Alaska (Lindroth 1963b: 378), south to northwestern Oregon (Westcott et al. 2006: 7), southern Colorado (LeConte 1879d: 501; Wickham 1902: 234; Elias 1987: 632), southwestern Nebraska (Keith County, CMNH), and southern South Carolina (Ciegler 2000: 49). The records from “Kansas” (Popenoe 1878: 79) and Arkansas (Wickham 1897: 104) need confirmation; those from Texas (Wickham 1897: 104), New Mexico (Fall and Cockerell 1907: 157) and Arizona (Wickham 1897: 104; Griffith 1900: 565) possibly refer to
Four North American species.
Bousquet and Webster (2006) provided a key for the identification of three species. Subsequently, a species described by Casey and incorrectly placed in another subgenus, as a synonym, was found to belong to this subgenus (see below under
This species is found from southern Alberta (Lindroth 1969a: 1115) to southeastern Arizona (Cochise County, CMNH), southern New Mexico (Hidalgo County, CMNH; Fall and Cockerell 1907: 157), and western Texas (Brewster and Culberson Counties, CMNH, MCZ).
This species is known from southern Saskatchewan, central South Dakota (Bousquet and Webster 2006: 32), and southeastern Colorado (Pueblo County, Foster F. Purrington pers. comm. 2009).
Members of this species are probably conspecific with
This species ranges from southern Manitoba to southern Alberta (Lindroth 1963b: 347), south to southern Texas (Kenedy County, CNC).
This species is known only from the holotype collected in southeastern Oregon.
This taxon has been listed as a junior synonym of
Northern Hemisphere, with eight species in the Nearctic (one Holarctic species) and Palaearctic (eight species) Regions.
Lindroth (1963b: 393-395) covered the North American species in his monograph of the Canadian and Alaskan
One species (
This Holarctic subspecies is found from northern Europe (Marggi et al. 2003: 248) to eastern Siberia (Kryzhanovskij et al. 1995: 82) and from the Chukchi Sea coast in Alaska (Lindroth 1963b: 394) to Newfoundland (Lindroth 1955a: 75-76), south to northern Pennsylvania (Bradford County, CMNH), northeastern Illinois (Lake County, CNC), east-central South Dakota (Kirk and Balsbaugh 1975: 20), and “Oregon” (Hatch 1953: 99).
The subspecies
Northern Hemisphere, with 26 species in the Nearctic (eight species) and Palaearctic (18 species) Regions.
Bousquet and Webster (2006) provided a key for the identification of all North American species except
This species inhabits the North American Cordilleras from the arctic circle in eastern Alaska and west-central Yukon Territory (Lindroth 1963b: 392) south to San Bernardino County in southeastern California (Dajoz 2007: 20, as
This species is known from Newfoundland to central Alaska, south to northeastern British Columbia and southern Alberta in the Rocky Mountains (Lindroth 1963b: 389). Fossil remnants of this species, dated between about 10,400 and 28,000 years B.P., have been unearthed in eastern Minnesota, northeastern Wisconsin, Illinois (Schwert 1992: 77), Iowa (Baker et al. 1986: 96; Schwert 1992: 77), central North Dakota (Ashworth and Schwert 1992: 260), and Cape Breton Island in Nova Scotia (Miller 1997: 250).
This eastern species ranges from Cape Breton Island to southern Saskatchewan (Lindroth 1963b: 388), south to eastern South Dakota (Larsen and Purrington 2010: 571), northern Illinois (Hayward 1897: 123), and northeastern West Virginia (Tucker County, CMNH).
This species ranges from New Brunswick and Maine west to eastern Minnesota and southeastern Manitoba (Bousquet and Webster 2006: 24).
This species is found from New Brunswick (Bousquet 1987a: 121) to the Osoyoos Valley in southern British Columbia, south to central Washington (Lindroth 1963b: 389), northern Iowa (Purrington et al. 2002: 201), northeastern Illinois (Lake County, CNC), and New York (Liebherr and Song 2002: 132; Saint Lawrence County, CMNH). The species seems to be local.
This species is found from the southern part of the Prairie Provinces (Lindroth 1963b: 393) to eastern Oregon (Baker County, CNC), south to Riverside County in southern California (Dajoz 2007: 19), central Nevada (Lander County, CNC), central Utah (Sevier County, CNC), southern Wyoming (Lindroth 1963b: 393), and southwestern North Dakota (Bowman County, CNC).
This species is known from Saskatchewan and Alberta (David R. Maddison pers. comm. 2012), southwestern Idaho (Owyhee County, CNC), southeastern Wyoming (Lavigne 1977: 44), and Costilla County in south-central Colorado (LeConte 1879d: 508; Wickham 1902: 234).
This eastern species ranges from Cape Breton Island (CNC) to “Michigan” (Garry A. Dunn pers. comm. 1986), south to northeastern West Virginia (Randolph County, CMNH).
About 90 species in the Nearctic (50 species), Neotropical (about 35 species), and Palaearctic (seven species) Regions. One species from the Philippines (
Lindroth (1963b: 357-375, as
This species is found along the Atlantic Coast from Long Island (Casey 1924: 41, as
This species is found along the Atlantic and Gulf of Mexico coasts from the Maritime Provinces (Lindroth 1963b: 373; Bousquet 1987a: 121) to southern Florida including the Keys (Leng 1915: 572), west to southeastern Texas (Wickham 1897: 104; Casey 1918: 124, as
This species is found along the Atlantic Coast from Newfoundland (Lindroth 1955a: 70), the Maritime Provinces (Lindroth 1963b: 373), and the Magdalen Islands (Larochelle 1975: 55) to southern Florida (Peck and Thomas 1998: 18) and along the Gulf Coast to southern Louisiana (Summers 1874a: 81; Hine 1906: 76; Allen 1965: 65). The records from “Ohio,” “Tennessee” (Hayward 1897: 109), eastern Kansas (Popenoe 1877: 24), and “Pennsylvania” (Bousquet and Larochelle 1993: 144) need confirmation.
This species is known so far only from the Florida Peninsula (Peck and Thomas 1998: 18).
This species is known from central Texas (Lee County, CMNH), southwestern Louisiana (Cameron Parish, Igor M. Sokolov pers. comm. 2009), and southwestern Florida (Monroe County, CMNH).
Hayward (1901: 158) listed
This species ranges from Connecticut (Krinsky and Oliver 2001: 88) to southeastern Alberta (Lindroth 1963b: 375), including southernmost Ontario (Bousquet 1987a: 121), north to Fort Smith in southern Northwest Territories (Lindroth 1963b: 374-375), south to northeastern Arizona (Apache County, CMNH), west-central New Mexico (Cibola County, CMNH), southern Texas (Johnson 1978: 67), and southern Florida, including the Keys (Peck and Thomas 1998: 18). The species is also known from the Bahamas (Darlington 1953: 5, as
This species is known only from the type locality, from Atlantic County in New Jersey (CMNH), and from one specimen simply labeled “Md” (CMNH).
This form was listed in synonymy with
This species is found from southeastern Manitoba to the foothills of the Rocky Mountains in southern Alberta, north to northeastern British Columbia (Lindroth 1963b: 359), south to northeastern Wyoming (Crook County, CNC), Kansas (Popenoe 1877: 24; Snow 1903: 193), Missouri (Summers 1873: 147; Lindroth and Freitag 1969: 337), southern Illinois (Hayward 1897: 101), and southwestern Ohio (Dury 1902: 111). The record from “Colorado” (Bousquet and Larochelle 1993: 141) needs confirmation.
The range of this species extends from New Brunswick to the foothills of the Rocky Mountains in Alberta (Lindroth 1963b: 375), south to south-central Montana (Carbon County, UASM), “Nebraska,” “Illinois” (Hayward 1897: 103, as
This species is found from Saskatchewan to western British Columbia, north to southern Northwest Territories (Lindroth 1963b: 361), south to central California along the coast (Motschulsky, 1859a: 131, as
This species ranges from New Brunswick (Larochelle and Larivière 1990a: 29) to northwestern Wisconsin (Hayward 1897: 100; Messer 2010: 36; Barron County, CMNH), south to northeastern Ohio (Lake and Trumbull Counties, CMNH, Harry J. Lee pers. comm. 2008) and northeastern West Virginia (Hampshire and Tucker Counties, CMNH). The record from southwestern Iowa (Wickham 1911b: 6) is probably in error.
This species has been recorded yet only from the type locality in southern Arizona.
The range of this species extends from southern Manitoba (Lindroth 1963b: 357) to eastern Washington (Hatch 1953: 96, as
This rarely collected species is known only from the Colorado River valley in southeastern California to western New Mexico (Cibola County, CMNH).
This species is known from southwestern California (Fall 1901a: 43; Moore 1937: 8).
This species ranges from the Okanagan Valley in southern British Columbia (Lindroth 1963b: 356) south to Inyo County in the Sierra Nevada of California (CMNH, MCZ, UASM) and southern Colorado (Elias 1987: 632). The record from southeastern South Dakota (Kirk and Balsbaugh 1975: 19) needs confirmation.
The range of this species extends from southwestern Manitoba to the Okanagan Valley in British Columbia (Lindroth 1963b: 356), south to central California along the Sierra Nevada (Dajoz 2007: 18), central Colorado (LeConte 1878a: 465; Wickham 1902: 234) along the Rocky Mountains, and northwestern Nebraska (Sheridan County, CNC).
This species is known from southeastern Manitoba to southwestern British Columbia (Lindroth 1963b: 365), south to eastern Washington (Hatch 1953: 96), northwestern Utah (Toole County, CMNH), south-central Colorado (Wickham 1902: 234; LeConte 1879d: 501; Lindroth 1963b: 365; Armin 1963: 165), central Nebraska (Blaine County, Foster F. Purrington pers. comm. 2010), and southeastern Wisconsin (Milwaukee County, Peter W. Messer pers. comm. 2009). The record from “Northwest Territories” (Bousquet and Larochelle 1993: 141) was based on four old specimens in the CNC labeled “N.W.T.” which does not refer to present day Northwest Territories or Nunavut.
This species is found from Skamania County in southern Washington (Hatch 1953: 93, as
This species is known only from the type series collected in central California.
The range of this species extends from western Newfoundland to the Rocky Mountains in Alberta (Lindroth 1963b: 367), south to north-central Colorado (Motz and Morgan 2001: Fig. 4d), northern Nebraska (Cherry County, Foster F. Purrington pers. comm. 2010), central Iowa (Story County, CNC), western West Virginia (Cabell County, CMNH), and Maryland (Erwin 1981b: 147). The records from northern Oklahoma (French et al. 2001: 228; Elliott et al. 2006: 125) need confirmation.
This species ranges from southern Manitoba to Vancouver Island (Lindroth 1963b: 358), north to the Great Slave Lake area in Northwest Territories (Bousquet 1987a: 121), south at least to Sierra County in the Sierra Nevada of California (MCZ; Lindroth 1963b: 358), the Grand Canyon in northern Arizona (CMNH), the Sacramento Mountains in southern New Mexico (CMNH), west-central Minnesota (Gandhi et al. 2005: 924), and central Wisconsin (Messer 2010: 36).
According to Erwin (1984a: 169), this species is found throughout California.
This species, as far as known, is restricted to the San Francisco Bay area in California (Erwin 1984a: 170).
This species is found from western New Hampshire (Cooper 1976: 164) to eastern North Dakota (Grand Forks County, CNC), north to southern Manitoba (Lindroth 1963b: 362) and southern Saskatchewan (Ronald R. Hooper pers. comm. 2002), south to southern Arizona (Snow 1906b: 161; Coconino County, CMNH), including north-central Utah (Cache County, CMNH), the Big Bend National Park in western Texas (Dajoz 2007: 23), west-central Arkansas (Garland County, Robert L. Davidson pers. comm. 2012), northern Tennessee (Montgomery County, Foster F. Purrington pers. comm. 2010), and west-central West Virginia (Roane County, CMNH).
This species is found along the Pacific Coast from southwestern British Columbia, including Vancouver Island (Lindroth 1963b: 369), south at least to west-central Oregon (Casey 1918: 104).
This species is known only from the type locality in southwestern Utah.
The range of this species extends from western Ontario (Lindroth 1963b: 369) to central Yukon Territory (Dawson, CNC), south to south-central Oregon (Lake County, UASM), southern Arizona (Casey 1918: 118, as
According to Erwin (1984a: 174), the range of this species “extends throughout the western United States in the mountainous regions;” the only documented records are those of the type localities in California and that from Valley County in central Idaho (Maddison 2012: Supplementary content Table S1). David R. Maddison (pers. comm. 2012) caught the species in Steens Mountains, southeastern Oregon.
This species is found from Kupreanof Island in the Alexander Archipelago to southern California (Moore 1937: 7; Kavanaugh 1992: 67).
This species ranges from southern Manitoba to southern British Columbia though not reaching the coast, north to Fort Smith in southern Northwest Territories (Lindroth 1963b: 363), south to northern Baja California (CNC), southern Arizona (Casey 1918: 119, as
This species occurs from the Saint Lawrence Plain in southern Quebec (Larochelle 1975: 57) to central British Columbia, north to east-central Alaska (Lindroth 1963b: 365), south to southern Colorado (Wickham 1902: 234; Bell 1971: 28), southern South Dakota (Kirk and Balsbaugh 1975: 18), southern Wisconsin (Messer 2010: 36), and southwestern Pennsylvania (Allegheny County, Robert L. Davidson pers. comm. 2008). The records from Missouri (Casey 1918: 109), southern Oklahoma (Hatch and Ortenburger (1930: 11), Texas (Wickham 1897: 104), Kansas (Popenoe 1877: 24; Snow 1878: 63), New Mexico (Fall and Cockerell 1907: 157), southern Arizona (Wickham 1897: 104), and “Delaware” (Bousquet and Larochelle 1993: 142) are probably in error.
This species is known only from the type locality in southwestern California.
This species is known only from the type series.
This species ranges from Newfoundland (Lindroth 1955a: 67) to Vancouver Island, north to central Alaska and the Mackenzie River delta in northern Northwest Territories (Lindroth 1963b: 370-371), south to El Dorado County in eastern California (Maddison 1985: 113), “Utah” (Hayward 1897: 105), northern New Mexico (Taos County, UASM), eastern North Dakota (Tinerella 2003: 635), northeastern Wisconsin (Messer 2010: 36), and the upper peninsula of Michigan (Casey 1924: 35, as
This species is yet known only from the lectotype.
This species is distinct from all other North American
This species is yet known only from the type locality and Santa Barbara County (Maddison 1985: 113) in western California.
This widely distributed species ranges from Newfoundland (Lindroth 1955a: 68) to western British Columbia, north to southern Northwest Territories (Lindroth 1963b: 372), south to Oregon (Hatch 1953: 94), southern Colorado (Elias 1987: 632), southern Texas (Kenedy County, CNC), northwestern Florida (Holmes County, UASM), and southern South Carolina (Ciegler 2000: 48).
This species is known only from the type locality in southeastern Arizona.
This species is known only from the California central valley (Erwin 1984a: 182).
This Holarctic species is found from Great Britain to eastern Siberia and Japan (Marggi et al. 2003: 254), and from the Arctic Plains in northern Alaska to the Hudson Bay shore in northwestern Ontario, south to southeastern Minnesota (Olmsted Country, CMNH), north-central Utah (Casey 1918: 105, as
This species is known from southern Texas (Johnson 1978: 67) and “Mexico” (Erwin 1984a: 185).
This species is known from the Gila River drainage in southwestern New Mexico (David R. Maddison pers. comm. 2007) and southern Arizona (Casey 1918: 116, as
This species is found from the Baja California Peninsula (Horn 1894: 308) and southeastern California (Casey 1918: 120; Andrews et al. 1979: 28) to southeastern Texas (Snow 1906a: 141; Casey, 1918: 121, as
The range of this species extends from Nova Scotia (Majka et al. 2007: 8) to the Rocky Mountains in Alberta (Lindroth 1963b: 364), south to southeastern Arizona (Dajoz 2007: 21; Cochise County, UASM), southern Texas (Johnson 1978: 67), northern Alabama (Madison County, CMNH), and eastern South Carolina (Ciegler 2000: 48), west along the southwest to southern California (Lindroth 1963b: 364; Los Angeles County, CMNH).
This species is known from the Gila River drainage in southwestern New Mexico (David R. Maddison pers. comm. 2007) and southeastern Arizona (Graham and Greenlee Counties, CMNH; Casey 1884b: 15, as
This taxon has been listed in synonymy with
This species is found from southern Manitoba to southern Alberta (Lindroth 1963b: 353), south to Mono County in east-central California (Dajoz 2007: 17), west-central Nevada (Bechtel et al. 1983: 474), central Colorado, and Nebraska (Hayward 1897: 88).
This species ranges from southern Manitoba to central British Columbia (Lindroth 1963b: 354, as
This species occurs from northwestern Ontario (CNC) to central Alberta (Lindroth 1963b: 355), south to central Utah (Fall 1922c: 171), southeastern Wyoming (Albany County, MCZ), and east-central South Dakota (Kirk and Balsbaugh 1975: 20). The record from “Colorado” (Bousquet and Larochelle 1993: 140) needs confirmation.
This species is found from southern Manitoba to the Okanagan Valley in British Columbia (Lindroth 1963b: 352), south to Kern County in southern California (Fall 1901a: 43), northern Utah (LeConte 1878a: 451; Casey, 1924: 40, as
Northern Hemisphere, with 16 species in the Nearctic (13 species) and Palaearctic (three species) Regions. The North American species are placed in two groups following Maddison (2012: 544).
Lindroth (1963b: 395-402) covered all North American species except for three (
The range of this species extends from southern Manitoba to Vancouver Island, north to east-central Alaska (Lindroth 1963b: 400), south to western Oregon (Hatch 1953: 100, as
This species is known from east-central Oregon (Union County, CMNH; Lindroth 1963b: 401) to southeastern California (Dajoz 2007: 20). The records from southwestern British Columbia and eastern Washington (Hatch 1953: 100) are probably in error (see Lindroth 1963b: 401).
This species is found from the Saint Lawrence Estuary in Quebec (Larochelle 1975: 53) to south-central British Columbia (Jarrett and Scudder 2001: 381) and central Washington (Grant County, Foster F. Purrington pers. comm. 2011), north to south-central Northwest Territories (near Hay River, CNC), south to southern Utah (Garfield County, FFPC).
This species is known from western California (Casey 1918: 160, 161, 163 as
This species is found from Vancouver Island (Lindroth 1963b: 401) to northwestern Montana (Russell 1968: 58), south to central New Mexico (Fall and Cockerell 1907: 157) and southwestern California (Moore 1937: 8).
This species is known from the type locality and Lincoln County in western Oregon (Maddison 2012: Supplementary content Table S1).
This name has been listed in synonymy with
This species is found from Cape Breton Island (Bousquet 1987d: 105) to south-central British Columbia, south to northern Idaho (Kootenai County, CNC), Missouri (Summers 1873: 147), and West Virginia (Pocahontas and Randolph Counties, CMNH). The records from “Washington,” “Oregon,” and “California” (Hayward 1897: 127, as
This species has passed under the name
This species is known only from the type locality in the Sierra Nevada.
This species is known from Vancouver Island (Lindroth 1963b: 402) to northwestern Montana (Russell 1968: 58), south to “Nevada” (Lindroth 1963b: 402) and Mendocino County in western California (Casey 1918: 157).
According to Lindroth (1963b: 396), this species is “probably restricted to C[entral] Calif[ornia].” It has been recorded so far from Placer (Casey 1918: 162, as
This species ranges from Newfoundland (Lindroth 1955a: 77, as
This species has passed under the name
This species occurs from Newfoundland (Lindroth 1963b: 397) to central Alaska (Lindroth 1963b: 397), south to northeastern Oregon (Umatilla County, MCZ), western Montana (Russell 1968: 57), western Nebraska (Keith County, CMNH), and northeastern West Virginia (Tucker County, CMNH). The records from central New Mexico (Fall and Cockerell 1907: 157, as
This species is found from Nova Scotia (Maddison 2012: Supplementary content Table S1) to the Okanagan Valley in British Columbia (Lindroth 1963b: 399), north to southern Northwest Territories (CNC), south to northern Illinois (Purrington et al. 2002: 200; McHenry County, MCZ), southwestern Pennsylvania (Allegheny County, CMNH), and Connecticut (Lindroth 1963b: 398).
Two North American species in the temperate regions.
Lindroth (1963b: 345-346) treated both species.
This species is confined to the west coast ranging from Vancouver Island (Lindroth 1963b: 345) to southern California (LeConte 1852a: 188; Fall 1901a: 42). The records from “Montana,” “Wyoming,” “Utah,” “Arizona” (Hayward 1897: 111), and “Colorado” (Csiki 1928: 118) almost certainly refer to
The range of this species extends from southern Manitoba (Lindroth 1963b: 346) to the Okanagan Valley in south-central British Columbia (Bousquet 1987a: 121), south at least to Inyo County in California (MCZ), “Arizona” (Lindroth 1963b: 346), southern Colorado (Alamosa County, CNC), and northeastern North Dakota (Ramsay County, MCZ).
Northern Hemisphere, with 21 species in the Nearctic (two species) and Palaearctic (19 species) Regions.
Lindroth (1963b: 381-382) treated both species found in North America.
This species ranges from central British Columbia to southwestern Manitoba (Lindroth 1963b: 382), south to northern Nebraska (Cherry County, Robert L. Davidson pers. comm. 2012), southwestern Colorado (Elias 1987: 632), and Mono County in east-central California (Dajoz 2007: 17).
This species is confined to the Pacific Coast from southern Vancouver Island (Lindroth 1963b: 382) to southern California (LeConte 1852a: 189; Fall 1901a: 43). The record from northern Idaho (Hatch 1953: 95) is probably in error.
Northern Hemisphere, with five species in the Nearctic (one Holarctic species) and Palaearctic (five species) Regions.
Lindroth (1963b: 297-298) covered the species found in North America.
This taxon is listed as a junior synonym of
The range of this Holarctic species extends from Norway to the Far East (Marggi et al. 2003: 246) in the Palaearctic Region and from the Seward Peninsula in Alaska east to northern Labrador (Lindroth 1963b: 298).
Northern Hemisphere, with about 75 species in the Nearctic (36 species) and Palaearctic (45 species) Regions. Only two species (
Lindroth (1963b: 268-297, as
The range of this species extends from “Nova Scotia” (Larochelle and Larivière 1990a: 28) to northeastern Ohio (Lee 1994: 58), south to western North Carolina (Swain County, CMNH) and south-central Tennessee (Grundy County, CMNH) along the Appalachian Mountains.
This species ranges from Newfoundland (Lindroth 1955a: 55) to southern Quebec (Larochelle 1975: 54), south to northeastern Tennessee (Greene and Unicoi Counties, CMNH) and northeastern Georgia (Fattig 1949: 18) along the Appalachian Mountains.
The range of this species extends from central Alaska to west-central Northwest Territories (Maddison 1985: 111, as
Maddison (2012: 535) noted that there at least two species within the current concept of this species.
This species is found along the Cascade Range and Coastal Ranges (Erwin 1984a: 172) from northwestern Washington to central California.
This subspecies ranges from southern Yukon Territory (Lindroth 1963b: 291) to northern Colorado along the Rocky Mountains (Casey 1918: 32) and to “California” (Lindroth 1963b: 291).
Three other subspecies of
This mountain species is known from southern British Columbia south at least to central Oregon along the Cascade Range (Lindroth 1963b: 286), west to Gallatin County in southern Montana (Maddison 2012: Supplementary content Table S1).
The range of this species extends from the Alaska Peninsula to southern Yukon Territory (Lindroth 1963b: 290), south at least to Ouray County in southwestern Colorado (CNC) along the Rocky Mountains (Lindroth 1963b: 289) and to the Sierra Nevada in eastern California (Dajoz 2007: 16).
This Holarctic species ranges from Scandinavia to eastern Siberia and the Far East (Marggi et al. 2003: 265) and from the Seward Peninsula in Alaska (Lindroth 1963b: 295) to the Hudson Bay in northeastern Manitoba (Garry 1993: 95).
This species is known only from a few localities in central and west-central British Columbia (Lindroth 1963b: 286).
This species is known from the Coast Ranges from Humboldt County (Notman 1929b: 222) to central California (Erwin 1984a: 178).
This species is known from the Kenai Peninsula in Alaska (Lindroth 1963b: 288) south to northern Washington (Glesne et al. 2000: 89) and central Alberta (Maddison 1985: 113). All records prior to Lindroth (1963b) listed by Bousquet and Larochelle (1993: 132) need confirmation.
The range of this species extends from the Haines district in southeastern Alaska to southeastern Yukon Territory (Lindroth 1963b: 287), south to western Montana (Russell 1968: 51) along the Rocky Mountains and foothills and to “California” (Lindroth 1963b: 287). The record from “Idaho” (Bousquet and Larochelle 1993: 132) needs confirmation.
This subspecies occurs from northern Alaska to southeastern Yukon Territory (Lindroth 1963b: 293), south to northwestern Montana (Edwards 1975: 54, as
This subspecies is found from Newfoundland (Lindroth 1955a: 54) to the Lake Superior area in western Ontario (Lindroth 1963b: 293), south to north-central Pennsylvania (Lycoming County, Robert L. Davidson pers. comm. 2008; Lindroth 1955a: 54).
This subspecies is reported from eastern Siberia and the Far East (Marggi et al. 2003: 266) but the specimens probably belong to a distinct (still undescribed) subspecies as pointed out by Lindroth (1963b: 293).
This species is known from a few mountain locations in the Similkameen district in southern British Columbia (Lindroth 1963b: 286) and from the Sierra Nevada in California (Casey 1924: 28; Maddison 1985: 114).
The range of this species extends from western Maine (Oxford County, MCZ; Larochelle and Larivière 1990a: 28) to southeastern Wisconsin (Rauterberg 1885: 22; Messer 2010: 36), including the Appalachian region in southern Quebec (Larochelle 1975: 62), south to Tennessee (Sevier County, CMNH) and North Carolina (Hayward 1897: 64; Brimley 1938: 117; Swain County, CMNH) along the Appalachian Mountains. The records from “Labrador,” “Hudson Bay Territory,” and “Missouri” (Hayward 1897: 64) refer to other species.
Lindroth (1963b: 268) included this species in its own group. However, I believe the species is closely related to some of the species included by Lindroth (1963b) in his
This species ranges from southwestern British Columbia, including Vancouver Island (Lindroth 1963b: 290), to southwestern Oregon (Hatch 1950: 98).
This subspecies is known from Alaska, including the arctic zone and Kodiak Island, southeastern Yukon Territory, and adjacent northern British Columbia (Lindroth 1963b: 294).
This subspecies ranges from Newfoundland (Lindroth 1955a: 52, as
This species is known from the California west coast region as far south as the San Francisco area (Casey 1918: 40, as
This Holarctic species ranges from western Siberia (Marggi et al. 2003: 265) to the Dempster Highway in Yukon Territory (Maddison 1985: 111; UASM), including the Aleutian Islands and Alaska Peninsula (Lindroth 1963b: 275), south to Saint Elias Mountains in northwestern British Columbia (Jarrett and Scudder 2001: 380).
This species is found from eastern Siberia (Marggi et al. 2003: 265) to the Hudson Bay coast in northern Manitoba (Holliday 1982: 116), including the Great Slave Lake area in Northwest Territories (Lindroth 1963b: 279).
The range of this species extends from the Aleutian Islands and the Gulf of Alaska coast to southwestern Yukon Territory (Lindroth 1963b: 273, as
The range of this species extends from the Aleutian Islands and the Alaska Peninsula (Lindroth 1963b: 283) south along the Rocky Mountains to northwestern Montana (Russell 1968: 51; Edwards 1975: 51) and to the central Sierra Nevada in California (Papp 1978: 164). The records from “Northwest Territories” (Bousquet and Larochelle 1993: 130) and from the San Juan River drainage in southwestern Colorado (Wickham 1902: 232) are probably in error.
This Holarctic species is found from eastern Siberia (Marggi et al. 2003: 265) to the Hudson Bay in northeastern Manitoba (Garry 1993: 95), including northern Alberta (Birch Mountains, Gerald J. Hilchie pers. comm. 2009). Fossil remnants of this species, dated between about 16,700 and 18,100 years B.P., have been unearthed in southeastern Iowa (Baker et al. 1986: 96).
This species is known from the Kenai Peninsula in Alaska (Lindroth 1963b: 278) south to “Oregon” (Hatch 1953: 84) and from north-central Colorado (Lindroth 1963b: 278).
This species ranges from British Columbia, as far north as the Prince Rupert area (Lindroth 1963b: 274), to Mendocino County in western California (Maddison 1985: 112) and to southern Colorado along the Rocky Mountains (Elias 1987: 632).
This species is known from Mount Rainier in Washington (Lindroth 1963b: 283) and the Sierra Nevada in California (Casey 1924: 25; Papp 1978: 164).
This species ranges from central Alaska including the Alaska Peninsula and Kodiak Island (Lindroth 1963b: 270) to western Northwest Territories (Tungsten, UASM), south at least to northern Utah (LeConte 1878a: 465, as
This species is known from Mount Rainier in Washington and the Sierra Nevada in California (Lindroth 1963b: 274). The record from British Columbia (Jarrett and Scudder 2001: 381) needs confirmation.
This species ranges from the Yukon River in western Alaska to the Mackenzie River delta in northwestern Northwest Territories (Lindroth 1963b: 278-279, as
This species is known yet only from the San Francisco Bay area in California.
This species is known from southeastern British Columbia (Syringa Provincial Park, James C. Bergdahl pers. comm. 2008) south at least to northern California (Casey 1924: 26, as
The range of this species extends from Newfoundland (Lindroth 1955a: 51) to southeastern Alberta (Lindroth 1963b: 282), south to east-central Minnesota (Aitkin County, CMNH) and mountains in New York and New England (Lindroth 1963b: 282). The four specimens labeled from the Anchorage area in southern Alaska (Lindroth 1963b: 282) could be mislabeled in my opinion. The record from “Pennsylvania” (Bousquet and Larochelle 1993: 131) needs confirmation.
According to Erwin (1984a: 179), this species is found in the foothills of the Sierra Nevada in California.
This species ranges from the Aleutian Islands and Alaska Peninsula (Lindroth 1963b: 276) to south-central Yukon Territory, south to the Sierra Nevada in California (Papp 1978: 164; Dajoz 2007: 16) and at least to northwestern Montana along the Rocky Mountains (Russell 1968: 50; Edwards 1975: 53).
This species is known from a few localities in northwestern Montana (Russell 1968: 51), southwestern Alberta, and southeastern British Columbia (Lindroth 1963b: 277).
This species is known from the Queen Charlotte Archipelago and adjacent mainland in British Columbia (Kavanaugh 1992: 65) and from northwestern Washington (Skagit County, James C. Bergdahl pers. comm. 2008).
Northern Hemisphere, with ten species in the Nearctic (four species) and Palaearctic (six species) Regions.
Lindroth (1963b: 250-254, as
Habu and Uéno (1955: 45) described the subgenus
This widely distributed species ranges from east-central Maine (Majka et al. 2011: 45) to “Montana” (Hayward 1897: 40) and southeastern Alberta (CNC), south to eastern Utah (Grand County, CNC), the Rio Grande in western Texas (Brewster County, CNC) and northern Chihuahua (Bates 1891a: 263), and the Florida Panhandle (Peck and Thomas 1998: 18).
This species occurs from the Laurentides region in western Quebec (Larochelle 1975: 57) to central Alaska (Lindroth 1963b: 253), south at least to northern Oregon (Morrow County, CMNH), northern Colorado (Eagle County, CMNH) along the Rocky Mountains, and southwestern North Dakota (Morton County, UASM).
The range of this species extends from Prince Edward Island (Majka et al. 2008: 130) and New Brunswick (Webster and Bousquet 2008: 16) to eastern Alaska (Lindroth 1969a: 1114), south to eastern Idaho (Clark County, UASM), southern Colorado (Wickham 1896c: 132; Maddison 1985: 95) along the Rocky Mountains, “Kansas” (Wickham 1896c: 132), Missouri (Summers 1873: 147), northwestern Indiana (Blatchley 1910: 69), and New York (Notman 1928: 214). One old specimen simply labeled “N.M.” is known (MCZ). The record from “New Jersey” (Wickham 1896c: 132) needs confirmation.
This species is known from scattered localities from northern Idaho (Bonner County, MCZ) to west-central Washington (Grays Harbor County, CMNH), south to southern California (Ventura County, CMNH). The record from British Columbia (Jarrett and Scudder 2001: 381) needs confirmation.
Northern Hemisphere, with seven species in the Nearctic (three species, of which two are adventive) and Palaearctic (six species) Regions.
Lindroth (1963b: 254-258, as
All recent authors seen (e.g., Marggi et al. 2003: 251; Lorenz 2005: 218; Ortuño and Toribio 2005: 278; Maddison 2012: 545) used
According to Maddison (2012: 569), this group is closely related to subgenus
This species ranges from southeastern Alaska to southern Yukon Territory (Lindroth 1963b: 256), south along the Rocky Mountains to central Colorado (Wickham 1902: 234; Hayward 1897: 115; Casey 1918: 115, as
This Palaearctic species is adventive in North America where it is known from southern British Columbia (Hatch 1953: 83; Smith et al. 2004: 96) to northwestern Oregon (Westcott et al. 2006: 7) in the west and from Newfoundland (Lindroth 1955a: 49; Larson and Langor 1982: 592) in the east. The first inventoried specimen collected in the eastern part of this continent was found in 1949 (see Lindroth 1955a: 49) and in the western part in 1947 (Hatch 1949c: 145). Fossil remnants from a Plio-Pleistocene sequence have been unearthed in northwestern Greenland (Böcher 1995: 24).
This Palaearctic species is adventive in North America where it is known from Newfoundland (Larson and Langor 1982: 592) to the coastal area of New Brunswick (Bousquet 1987a: 120), including the Magdalen Islands in the Gulf of Saint Lawrence (Larochelle 1975: 60), and from southeastern Maine (Larochelle and Larivière 1990a: 28, 33). One specimen has been collected recently in a campground in Kittitas County, central Washington (Robert L. Davidson pers. comm. 2008). The first inventoried specimen collected on this continent was found in Nova Scotia in 1942 (Majka et al. 2007: 7).
One species in western North America.
The species is described in detail, under the name
This species is known from the Cascade Range in southwestern Oregon south to central California along the Coast Ranges (Casey, 1918: 44, as
Eleven species in the Nearctic (five species) and Palaearctic (seven species) Regions. One species (
Bousquet and Webster (2006) provided a key for the identification of all North American species. Lindroth (1963b: 348-351, 360) covered four species, three he placed in the
The range of this species extends from Nova Scotia (CNC) to eastern Minnesota (Epstein and Kulman 1990: 214), south to east-central Iowa (Johnson County, USNM), west-central Indiana (Blatchley 1910: 76), and southeastern West Virginia (Greenbrier County, MCZ). The record from southern South Dakota (Kirk and Balsbaugh 1975: 19) needs confirmation.
This species occurs from Newfoundland (Lindroth 1955a: 66) to southeastern Ontario (Ottawa, CNC), south to New England and the Adirondack Mountains in northeastern New York (Lindroth 1963b: 349). The record from eastern Iowa (Cooper 1976: 163) needs confirmation.
This species ranges from Newfoundland (Lindroth 1955a: 64) to east-central Alaska (Lindroth 1963b: 349), south at least to Sonoma County in western California (Casey, 1918: 96, as
This species has passed for a long time under the name
This species ranges from New Brunswick to southwestern Quebec, south to Virginia (Bousquet and Webster 2006: 29).
The range of this species extends from New Brunswick (CNC) to eastern South Dakota (Kirk and Balsbaugh 1975: 19), south to western (Dajoz 2007: 23) and east-central (Tucker 1906: 85) Texas, southwestern Alabama (Baldwin County, CMNH), northeastern Georgia (Leng 1910: 73; Fattig 1949: 17), and eastern South Carolina (Ciegler 2000: 49). The record from Colorado (Wickham 1902: 234) needs confirmation; that from British Columbia (Jarrett and Scudder 2001: 382) was based on misidentified specimens of
This species has been listed in the subgenus
Western Nearctic Region, with two species.
Both species are included in Lindroth’s (1963b: 266-267) monograph.
This species ranges from central Alberta (Lindroth 1963b: 267) to the Queen Charlotte Islands (Kavanaugh 1992: 64), south to southern California (Fall 1901a: 10; Moore 1937: 6) and northern Utah (Knowlton 1939: 2; Davis and Salt Lake Counties, CMNH).
This species is known from southern California (Fall 1901a: 43; Moore 1937: 6; Maddison 2012: Supplementary content Table S1). The record from “Washington” (Hayward 1897: 128) was based on a misidentified
Three western North American species.
Lindroth (1963b: 343-345) treated two species (
This species has been recorded from Benton and Jackson Counties in western Oregon (Westcott et al. 2006: 7), from Humboldt and Yolo Counties in northern California (Maddison 2012: Supplementary content Table S1), and from southwestern California (Casey 1918: 45; Moore 1937: 6).
This species is known from southern California (Fall 1901a: 43; Moore 1937: 6), northern Arizona (Navajo and Apache Counties, UASM; Hayward 1897: 117), southern Utah (San Juan and Kane Counties, MCZ, UASM), and western New Mexico (Maddison 2012: Supplementary content Table S1). The records from “Washington” and “Oregon” (Bousquet and Larochelle 1993: 139) are likely in error.
This species is found from southwestern British Columbia, including Vancouver Island (Lindroth 1963b: 344), south to southern California (Fall 1901a: 43, as
Northern Hemisphere, with two species in the Nearctic (
The North American species is treated in Lindroth’s (1963b: 284) monograph.
This subgenus was placed in synonymy with
This species ranges from Newfoundland (Lindroth 1955a: 52) to the Norton Sound area in Alaska (Lindroth 1963b: 284), south to central California along the Coast Ranges, southern Colorado along the Rocky Mountains, Isle Royale in northernmost Michigan (Hubbard and Schwarz 1878: 629), and Cape Breton Island [see Lindroth 1963a: Fig. 63]. Fossil remnants from a Plio-Pleistocene sequence have been unearthed in northwestern Greenland (Böcher 1995: 24).
Northern Hemisphere, with 22 species in the Nearctic (five eastern species) and Palaearctic (17 Asian species) Regions (see Toledano and Schmidt 2010).
Four of the North American species are included in Lindroth’s (1963b: 212-229) key to species of
This eastern species ranges from “Vermont” (Hayward 1897: 72) and southwestern Massachusetts (Hampden County, MCZ) to “Illinois” (Hayward 1897: 72), south to northern and eastern Tennessee (Cheatham, Fentress, and Sevier Counties, CMNH, MCZ). The record from southern Wisconsin (Rauterberg 1885: 22) needs confirmation; those from Missouri (Summers 1873: 147) and the Rocky Mountains in northern Colorado (Wickham 1902: 232) must be in error.
This species is known from central Iowa (Maddison 2012: Supplementary content Table S1), northeastern Kansas (Pottawatomie County, CMNH; Lindroth 1963b: 218; Knaus 1903: 188), eastern Oklahoma (Latimer County, UASM) and central Texas (Blanco County, UASM; Hayward 1897: 70). According to Hildebrandt and Maddison (2011: 273), the records from Massachusetts, “New Jersey,” “Pennsylvania,” the District of Columbia (Hayward 1897: 70), and New York (Notman 1928: 216) are likely incorrect and may be based on
This species is known from the Ozark Plateau of Missouri and Arkansas [see Hildebrandt and Maddison 2011: Fig. 6].
The range of this eastern species extends from Nova Scotia (Lindroth 1954c: 302) to “Minnesota” (Hildebrandt and Maddison 2011: 274), south to eastern Oklahoma (Le Flore County, FFPC ), northern Arkansas (Lindroth 1969a: 1114; Pope, Searcy, and Washington Counties, CMNH), northern Mississippi (Drew A. Hildebrandt pers. comm. 2010), central Alabama (Jefferson County, Drew A. Hildebrandt pers. comm. 2009), northern Georgia (Fattig 1949: 17), and northwestern South Carolina (Ciegler 2000: 48).
This eastern species ranges from Nova Scotia (Larochelle and Larivière 1990a: 28; Maddison 2012: Supplementary content Table S1) to the Ontario Peninsula (Lindroth 1963b: 300) and northern Ohio (Ashtabula, Cuyahoga, Erie, and Lorain Counties, Harry J. Lee, Jr. pers. comm. 2008), south to west-central Tennessee (Hickman County, CMNH) and northeastern Virginia (Richard L. Hoffman pers. comm. 1992).
West Palaearctic Region, with nine species of which one is adventive in eastern North America.
The species found in North America was covered in Lindroth’s (1963b: 258) monograph.
Maddison (2012: 561) noted that this taxon may be the sister-group of the remaining
This Palaearctic species is adventive in North America where it is known from Prince Edward Island (Majka et al. 2008: 130), and from western Quebec, as far north as the Abitibi region (Paquin and Dupérré 2002: 86), to the Algoma District in northern Ontario (Pearce et al. 2003), south to southern Michigan (Purrington et al. 2002: 200), northeastern Ohio, and Long Island, New York [see Hoebeke et al. 1991: Fig. 1]. The first inventoried specimen collected on this continent was found in the Great Lake region in Ontario in 1956 (Lindroth 1963b: 258).
Northern Hemisphere, with four coastal species in North America (two species, one endemic along the Pacific and one adventive species along the Atlantic) and Europe (three species:
Both species found in North America are included in Lindroth’s (1963b: 213) key to the Canadian and Alaskan
This species occurs on the seashore along the coast of California.
This adventive species was known on this continent until recently only from the four specimens upon which Hayward (1897: 129) described his
One species in western North America.
Lindroth (1963b: 407-408) covered the species in his monograph of the Canadian and Alaskan
Molecular data analyses presented by Maddison and Ober (2011) and Maddison (2012) strongly suggest that this genus is not closely related to the other members of
This species ranges from Vancouver Island (Lindroth 1963b: 408) to western Montana (Russell 1968: 58), south to Nevada County in the Sierra Nevada (Dajoz 2007: 16) and to Sonoma County along the Pacific Coast (Lindroth 1963b: 408).
New World, with about 125 species arrayed in seven genera:
Western Hemisphere, with 13 species in temperate, subtropical, and tropical areas of the Nearctic (one species) and Neotropical (12 species) Regions.
The North American species is treated in Lindroth’s (1966: 441) monograph under the genus
This species ranges from the Nova Scotia Peninsula (Christopher G. Majka pers. comm. 2007) to western Washington (Hatch 1953: 104), north to southern British Columbia (Lindroth 1966: 441) and southern Alberta (CNC), south to southwestern California (Los Angeles County, CAS), southeastern Arizona (Dajoz 2007: 21), south-central Texas (Bastrop County, CNC), and southern Florida (Peck and Thomas 1998: 18).
Worldwide, with about 790 species (Lorenz 2005: 207-215). The Northern Hemisphere is represented by about 230 species (roughly 29% of the world fauna) and North America by 73 species (9.2% of the world fauna). Two species are adventive (
The genitive form of
Worldwide, with 24 species in boreal, temperate, subtropical, and tropical areas. The species are arrayed in two subgenera:
Erwin (1975) revised the species and provided a key for their identification.
Worldwide, with 19 species in the Nearctic (five species), Neotropical (three species, only one of them endemic), Australian (five species), Oriental (four species), Palaearctic (two species), and Afrotropical (four species) Regions.
This species ranges from Cape Breton Island (Lindroth 1954c: 303) to west-central Yukon Territory, south to Veracruz in Mexico, northeastern Mississippi (Oktibbeha County, Drew A. Hildebrandt pers. comm. 2009), and northern Georgia [see Erwin 1975: Fig. 121]. The record from “Oregon” (Bousquet and Larochelle 1993: 149) needs confirmation.
This western species ranges from southwestern British Columbia, including Vancouver Island, to western Idaho, south to Fresno County in California along the Sierra Nevada and to the San Francisco Bay area along the Coast Ranges [see Erwin 1975: Fig. 111].
This species ranges from Maine and southern Quebec to south-central British Columbia, south to southwestern California, Belize and Guatemala, and southern Florida, including the Keys; also known from Cuba [see Erwin 1975: Fig. 135].
The range of this species extends from the Gaspé Peninsula in eastern Quebec to west-central British Columbia, south to northwestern Oregon, south-central New Mexico through the Rocky Mountains, the Black Hills in southwestern South Dakota, east-central Minnesota, western New York near the Great Lakes, and Massachusetts (Erwin 1975: 65) along the east coast [see Erwin 1975: Fig. 135]. The record of
This species ranges from Massachusetts to the Black Hills in southwestern South Dakota, south to southern Arizona, southern Texas, southern Mississippi (Harrison and Lamar Counties, Drew A. Hildebrandt pers. comm. 2008), and southern Florida, including the Keys [see Erwin 1975: Fig. 156].
Worldwide, with about 340 species (Lorenz 2005: 207-211, as
Several authors have discussed the taxonomic status of
Western Hemisphere, with 37 species in North America (28 species), Middle America (ten species), and the West Indies (three species, two of them endemic).
Hayward (1900: 201-212, as
Sciaky and Vigna Taglianti (2003) considered
The range of this species extends from Cape Breton Island to southeastern Alberta, north to southern Northwest Territories (Lindroth 1966: 418-419), south to southeastern Arizona (Dajoz 2007: 21), northwestern Texas (Nolan County, CMNH), northwestern Louisiana (Caddo Parish, Igor M. Sokolov pers. comm. 2009), and Maryland (Clark et al. 2006: 1306). The records from “British Columbia,” “Washington,” “Oregon,” “Idaho,” “Utah” (Bousquet and Larochelle 1993: 150), based on specimens identified under this name in USNM, need confirmation.
Lindroth (1966: 417) considered
This species ranges from southern Vancouver Island (CNC) to northern Idaho (Hatch 1953: 103), south to southern California (LeConte 1852a: 192; Fall 1901a: 44).
This species is known only from the type locality in southeastern Virginia.
This form was considered a synonym of
This species is currently known only from the Coastal Plain ranging from New Jersey (Casey 1884c: 71, as
The range of this species extends from western Vermont (Addison and Chittenden Counties, CMNH) to eastern Iowa (Hayward 1900: 209), including southernmost Ontario (Bousquet 1987a: 122), south to southeastern Louisiana (East Baton Rouge and Saint Tammany Parishes, Igor M. Sokolov pers. comm. 2009), southern Mississippi (Peter W. Messer pers. comm. 2008), and southern Florida (Peck and Thomas 1998: 18).
This form was regarded as a valid species by Hayward (1900: 209), Casey (1918: 187), Lindroth (1966: 422), and Bousquet (1987a: 122) but listed as a synonymy of
This species is known from northeastern New Mexico (Fall 1907: 218) and southeastern Arizona (Cochise County, USNM).
This species has been reported from localities in South Dakota (Kirk and Balsbaugh 1975: 21), west-central Mississippi, and eastern and central Texas (Casey 1918: 181).
This form was considered a synonym of
This species is known only from southern Arizona.
This species ranges from southern New Hampshire (Hillsborough County, CMNH) to south-central South Dakota (Kirk and Balsbaugh 1975: 21), including southern Ontario (Lindroth 1966: 419), south to southern Texas (Casey 1918: 177, as
This species is known only from the type locality in southern Florida.
This species ranges from “Massachusetts” (Hayward 1900: 212) and Connecticut (Krinsky and Oliver 2001: 90) to northeastern Kansas (Popenoe 1878: 79), including eastern Iowa (Lindroth 1966: 423), south to northwestern and central Texas (Barr 1964: 3), southern Alabama (Conecuh County, CMN), and the Florida Panhandle (Washington County, CNC). The records from “Colorado” (Beutenmüller 1897: 39) and “New Hampshire” (Bousquet and Larochelle 1993: 151) need confirmation.
Under this name, LeConte (1863b: 15) listed “
This species seems restricted to the Coastal Plain ranging from New Jersey (Smith 1890: 79; Smith 1910: 203) to southern Florida (Leng 1915: 573), west to “Louisiana” (Chaudoir 1868b: 214) including southwestern Alabama (Löding 1945: 14).
The range of this species extends from Nova Scotia (Lindroth 1966: 416) to “Nebraska” (Hayward 1900: 204), south to northern Oklahoma (French et al. 2001: 228), southeastern Louisiana (East Baton Rouge, Tangipahoa, and Saint Tammany Parishes, Igor M. Sokolov pers. comm. 2009), and northern Florida (Peck and Thomas 1998: 18). The records from southern Colorado (Elias 1987: 632) and “Minnesota” (Bousquet and Larochelle 1993: 151) need confirmation.
This species ranges from Cape Breton Island (Bousquet 1987a: 121) to the Okanagan Valley in south-central British Columbia (Lindroth 1966: 416), south to “Oregon” (Hayward 1900: 205; Hatch 1953: 104), central New Mexico (Fall and Cockerell 1907: 157), and southeastern Tennessee (Sequatchie and Bledsoe Counties, Robert L. Davidson pers. comm. 2009). Several state records listed in Bousquet and Larochelle (1993: 151) need confirmation.
This species is known only from the type series.
This species is known only from the type locality in northwestern New Mexico.
This species is known from North Carolina (Brimley 1938: 118) to “Utah” (Hayward 1900: 207), south to southeastern Arizona (Cochise County, USNM), southeastern Texas (Casey 1918: 178, as
This form was listed as a synonym of
This species is known from Sonora in Mexico (USNM), southern Arizona (Pima and Santa Cruz Counties, CMNH, USNM), and “California” (Hayward 1900: 211). The record from southwestern Utah (Tanner 1928: 270) needs confirmation.
This species is known from southeastern Washington (Walla Walla County, MCZ) and northern Idaho (Hatch 1953: 103) south to southern California (Fall 1901a: 43) and central Arizona (Griffith 1900: 565; Hayward 1900: 206; Casey 1918: 177), east along the south to western Texas (Casey 1918: 177; El Paso County, MCZ).
This form has been listed in synonymy with
This species is known from east-central Oregon (Grant County, James R. LaBonte pers. comm. 1992), Humboldt County in northwestern California, and Washoe County in northwestern Nevada (Casey 1918: 183).
This species is known from “Indiana” to southeastern Iowa (Casey 1918: 186), south to the Rio Grande in southern Texas (Casey 1918: 188, as
The range of this species extends from Nova Scotia (Lindroth 1966: 422) to southwestern Wisconsin (Messer 2010: 37), south to central Louisiana (Grant Parish, CMNH), southwestern Alabama (Clarke County, CMNH), and northern Georgia (Fattig 1949: 19). The records from “Minnesota” and “Kansas” (Bousquet and Larochelle 1993: 152) need confirmation.
This species is known from “Arizona” and northern Utah (Casey 1918: 180).
This species has been reported from northwestern Nevada (Casey 1918: 185, as
This species is known only from the type locality in the Sierra Nevada.
This species ranges from Cape Breton Island (Lindroth 1966: 423) to “Iowa” (Hayward 1900: 210), south to southern Texas (Casey 1918: 189, as
This species is known from Nova Scotia (Majka et al. 2007: 8) to southern Saskatchewan (Ronald R. Hooper pers. comm. 2008), south to Arkansas (Franklin County, CNC) and North Carolina (Casey 1918: 183, as
The range of this species extends from Nova Scotia (Majka and Vickery 2008) to “Wisconsin” (Rauterberg 1885: 23), including southern Quebec and southern Ontario (Lindroth 1966: 423), south to northwestern Texas (Nolan County, CMNH), southwestern Alabama (Clarke County, CMNH), and northern Florida (Suwanee County, CMNH). The record from southern Arizona (Snow 1907: 142) is probably in error.
This species ranges from Nova Scotia (Lindroth 1966: 422) to western Kansas (Trego County, CNC), south to southeastern Texas (Wickham 1897: 104; Cameron County, CNC) and central Florida (Peck and Thomas 1998: 18). The species was also recorded from the Bahamas (Darlington 1953: 5) and Cuba (Darlington 1934: 78).
According to Sciaky and Vigna Taglianti (2003: 90), members of
The species was included in Lindroth’s (1966: 410) key to the species of
This European species is adventive in North America where it is known from southwestern British Columbia, Washington, Oregon, and Idaho [see LaBonte and Nelson 1998: Fig. 1]. The first inventoried specimen reported from this continent was collected in Seattle in 1940 (Hatch 1950: 105).
Western Hemisphere, with five species in the Nearctic (one species) and Neotropical (four species) Regions, including the West Indies. Erwin (1974a: 125) stated that he was aware of 35 undescribed species in this genus.
Barr (1971b) redescribed the North American species and illustrated its male genitalia.
This taxon is possibly the sister-group to
This species occurs from Connecticut (Krinsky and Oliver 2001: 96) to “Indiana” (Schrock 1985: 351), south to southeastern Texas (San Patricio County, UASM) and southern Florida including the Keys (Peck and Thomas 1998: 18), west along the southwest to southwestern California (Riverside County, CAS); one specimen, possibly mislabeled, was seen from western Oregon (Benton County, CNC). The record from “Illinois” (Bousquet and Larochelle 1993: 152) needs confirmation.
New World, with about 70 species arrayed in three subgenera:
Erwin (1974b) revised the species and provided a key for their identification.
Western Hemisphere, with about 45 species in the Nearctic (three species) and Neotropical (about 45 species) Regions.
This species ranges from southwestern New Hampshire (Choate 1977: 115) to northern Kansas, south to Honduras and central Florida [see Erwin 1974b: Fig. 73]. The record from southern Wisconsin (Rauterberg 1885: 23) needs confirmation.
This species is known from central Oklahoma (Grady County, CMNH), southwestern New Mexico, and southeastern Arizona, south to Durango in Mexico. The species is apparently absent from the desert region of northern Mexico [see Erwin 1974b: Fig. 75].
This species is known only from “California” and southern Arizona [see Erwin 1974b: Fig. 134].
Five species in the Palaearctic (three species) and Oriental (two species) Regions. Among the Palaearctic species, one is endemic to Japan (
The species found in North America was covered in Lindroth’s (1966: 424) monograph.
This Palaearctic species is adventive in North America where it ranges from Nova Scotia (Majka et al. 2006: 605) to eastern North Dakota (Cass County, Donald P. Schwert pers. comm. 1989), south along the east coast to New Jersey (Smith 1910: 203, as
Worldwide, with about 40 species in the Nearctic (three species), Neotropical (ten species), Australian (nine species), Oriental (eight species), Palaearctic (nine species), and Afrotropical (ten species) Regions. One species is also known from the Hawaiian Islands.
Lindroth (1966: 424-427) treated two (
This taxon is considered a subgenus of
This species is known only from the type locality in central Texas.
This species ranges from Nova Scotia (Bousquet 1987a: 123) to southwestern North Dakota (Smith et al. 1979: 92), south to southeastern Texas (Casey 1918: 213) and southern Florida including the Keys (Peck and Thomas 1998: 18).
This species is known from central Idaho (Custer County, Ken Karns pers. comm. 2009), southeastern Oregon (Malheur County, James R. LaBonte pers. comm. 1992), California, from Lake County (CAS) to Riverside County (CAS) east to Mono and Inyo Counties (CAS; Dajoz 2007: 18), southern Arizona (Cochise and Maricopa Counties, CMNH; Hayward 1900: 217), southern Texas (Zapata County, CMNH; Casey 1918: 211), northwestern Arkansas (Newton County, Peter W. Messer pers. comm. 2008), and eastern South Dakota (Kirk and Balsbaugh 1975: 21; French et al. 2004: 557). The record from Aklavik in northern Northwest Territories (Lindroth 1966: 427) is quite obviously based on a mislabeled specimen.
Worldwide, with about 180 species (Lorenz 2005: 212-214, as
The first use of the name
Shilenkov (2002) treated
About 35 species in the Nearctic (11 species), Neotropical (seven species), Australian (one non-endemic species), Oriental (five species), Palaearctic (12 species, none in the Far East), and Afrotropical (seven species, three shared with Eurasia) Regions. One species,
Lindroth (1966: 427-431) treated five (one of them,
This species has been reported from a few islands in the West Indies (Peck and Thomas 1998: 18; Steiner 2008: 131), “Louisiana” (Hayward 1900: 224), and southeastern Texas (Casey 1918: 208); it is also known from southern Mississippi (Hancock and Jackson Counties, Drew A. Hildebrandt pers. comm. 2009) and Grady County in central Oklahoma (Robert L. Davidson pers. comm. 2012). The record from “Florida” (Bousquet and Larochelle 1993: 153) needs confirmation.
This species is still known only from the holotype.
This species ranges from eastern Washington (Hatch 1953: 104) to “Montana” (Hayward 1900: 223), south to western Texas (Dajoz 2007: 23), southeastern New Mexico (Chaves County, CNC), and the Baja California Peninsula (Horn 1894: 308). The records from “Nebraska” and “Texas” (Hayward 1900: 223) probably refer to
This species is known from the southern parts of the Prairie Provinces (Lindroth 1966: 429; Bousquet 1987a: 123) and eastern North Dakota (Grand Forks County, UASM, USNM). The records of
This species is found along the Coastal Plain from Long Island, New York (Cooper 1935: 144) to southern Florida, including the Keys (Peck and Thomas 1998: 18), west to southeastern Texas (Casey 1918: 207, as
This species has been reported from southwestern Alabama (Löding 1945: 14), “Texas” (LaFerté-Sénectère 1841a: 48), and Guatemala (Bates 1882a: 139, as
This species has been recorded from “California,” “Arizona,” and Salt Lake Valley in Utah (Hayward 1900: 220). The record from “Colorado” (LeConte 1858a: 28) probably refers to the Colorado River.
This species is known from New Jersey (Beutenmüller 1897: 37; Hayward 1900: 221; Smith 1910: 203), the Florida Peninsula (Leng 1915: 574), Big Pine Key, one of the Florida Keys (Foster F. Purrington pers. comm. 2012), Claiborne County in southwestern Mississippi (Robert L. Davidson pers. comm. 2012), Cameron Parish in southwestern Louisiana (Hine 1906: 76), “several localities” in Texas (Hayward 1900: 221), and Chaves County in southeastern New Mexico (Robert L. Davidson pers. comm. 2012); it has been recorded also from Guana Island (Valentine and Ivie 2005: 275) in the British Virgin Islands.
This species is known from southeastern Georgia (Glynn County, CMNH) and the Florida Peninsula (Monroe and Pinellas Counties, CMNH; Hayward 1900: 221, as
This species is known from Cuba and from Big Pine Key, Monroe County, in southern Florida (Foster F. Purrington and Robert L. Davidson pers. comm. 2009). This is a
This species is found along the seacoast from Santa Barbara County (Fall 1901a: 43) to San Diego County (LeConte 1852a: 194; Moore 1937: 8) in southwestern California.
This species is known from Vancouver Island (Lindroth 1966: 428, listed as “doubtful”), along the seacoast in southern California, including Santa Catalina Island (LeConte 1852a: 193; Fall 1901a: 43), Baja California Norte (Moore and Legner 1974: 289), Guatemala (Bates 1882a: 139), the Galápagos Islands (Mutchler 1925: 223, as
Worldwide, with about 145 species in the Nearctic (20 species), Neotropical (about 35 species), Australian (about 15 species), Oriental, Palaearctic (about 30 species), and Afrotropical Regions. Erwin (1974a: 126) stated that the number of New World species known to him is over 300.
Lindroth (1966: 431-436) treated six and Hayward (1900: 224-232) 11 (ten in the key) of the 20 species found in North America. A revision of the group is needed.
This species has been reported from southern Alabama (Motschulsky 1862b: 29) and western Mississippi (Casey 1918: 201).
This species is known from the Coastal Plain ranging from North Carolina (Brimley 1938: 118) to southern Florida (Peck and Thomas 1998: 18), west to “Louisiana” (LeConte 1863c: 20), and from Cuba (Darlington 1934: 80) and Guadeloupe (Fleutiaux and Sallé, 1890: 368, as
This species is known from southwestern Pennsylvania (Allegheny County, CMNH), Virginia (Hoffman et al. 2006: 21), South Carolina (Ciegler 2000: 54), east-central Georgia (Emanuel County, Robert L. Davidson pers. comm. 2012), central Florida (Pinellas County, CMNH), southwestern Alabama (Baldwin County, Robert L. Davidson pers. comm. 2012), northeastern Mississippi (Oktibbeha County, Drew A. Hildebrandt pers. comm. 2009), Texas (Cameron, Nolan, Travis, and Zapata Counties, CMNH; Riley 2011), and central Arkansas (Pulaski County, CMNH). The record from South Dakota (Kirk and Balsbaugh 1975: 21) needs confirmation.
This species seems confined to the Coastal Plain and Piedmont Plateau ranging from southeastern Pennsylvania (Lebanon and Dauphin Counties, CMNH) to southern Florida, including the Keys (Peck and Thomas 1998: 18), west to Arkansas (Pulaski and Garland Counties, CMNH) and eastern Texas (Sabine County, Robert L. Davidson pers. comm. 2012), including Alabama (Löding 1945: 14) and Winston County in east-central Mississippi (Drew A. Hildebrandt pers. comm. 2009).
This species ranges from south-central British Columbia (Lindroth 1966: 436) south to “Utah” (Hayward 1900: 231) and at least Santa Clara in west-central California (Erwin 1974a: 140).
This species has been reported only from central Texas (Casey 1918: 200) and southeastern New York (Casey 1918: 200, as
This species ranges from southern Quebec (Larochelle 1975: 111, as
This species is known from south-central Ohio (Fairfield County, CMNH) to “Massachusetts,” south to “North Carolina” (Erwin 1981b: 152).
This species ranges from southeastern Maine (Majka et al. 2011: 46) and the Saint Lawrence Plain in southern Quebec (Larochelle 1975: 111) to eastern South Dakota (Kirk and Balsbaugh 1975: 21), south to southeastern New Mexico, northern Coahuila in Mexico (Barr and Reddell 1967: 269), and central Florida (Seminole County, CMNH). The record from Jamaica (Darlington 1941a: 11) needs confirmation.
The range of this species extends from southwestern Quebec (LeSage 1996: 23) to southern Wisconsin (Messer 2010: 37), south to southeastern Texas (Kleberg County, CMNH; Hayward 1900: 230) and southern Florida (Peck and Thomas 1998: 18); also recorded from the Bahamas (Darlington 1953: 6, as
This species is known only from the holotype.
None.
This species is known from scattered localities from New Brunswick (Webster and Bousquet 2008: 17) to northern Ohio (Cuyahoga County, Harry J. Lee, Jr. pers. comm. 2008), south to southern South Carolina (Ciegler 2003: [2]) and southwestern Alabama (Baldwin County, CMNH).
Besides the type locality, this species is known from several counties in Virginia (Hoffman et al. 2006: 21), Maryland (Lepping 2009: 65), Montgomery County in northern Tennessee (Foster F. Purrington pers. comm. 2011), Oktibbeha County in northeastern Mississippi (Drew A. Hildebrandt pers. comm. 2009), and Desha County in southeastern Arkansas (Foster F. Purrington pers. comm. 2010).
This species ranges from New Brunswick (Bousquet 1987a: 124) to central South Dakota (Kirk and Balsbaugh 1975: 21), south to central Texas (Blanco and Travis Counties, CMNH; Hayward 1900: 229; Riley 2011) and southern Florida (Peck and Thomas 1998: 18); also recorded from Cuba (Darlington 1934: 81) and Jamaica (Darlington 1941a: 12).
This species is known only from the lectotype. The record from “Colorado” (Leng 1920: 54) needs confirmation.
This species is known only from east-central and westernmost Texas (Casey 1918: 202).
This species has been reported from “Arkansas” (Casey 1918: 193), “Louisiana” (Chaudoir 1868b: 213), and southeastern Texas along the Gulf Coast (Casey 1918: 193).
This form has been listed as a synonym of
This species is known along the Coastal Plain from southwestern Georgia (Fattig 1949: 19) to southern Florida (Peck and Thomas 1998: 18), west to southeastern Texas (Casey 1918: 197).
This species is known only from the type locality in southeastern Texas.
This species is found from southeastern Texas (Wickham 1897: 104) to southwestern California (Fall 1901a: 43; Moore 1937: 8) and the Baja California Peninsula (Horn 1894: 308), north to southwestern Utah (Tanner 1928: 270). The records from the Bahamas (Darlington 1953: 6), Cuba (Darlington 1934: 82; Mateu 1977: 378), Jamaica (Darlington 1941a: 11), and Puerto Rico (Wolcott 1936: 188) need confirmation; that from eastern Washington (Hatch 1953: 105) is probably in error; that from “Colorado” (LeConte 1858a: 28) probably refers to the Colorado River.
Worldwide, with about 375 species in the Nearctic (50 species), Neotropical (24 species), Australian (about 45 species), Oriental (about 22 species), Palaearctic (about 175 species), and Afrotropical (about 60 species) Regions. There is no genus-group taxa shared between the Western Hemisphere and the Eastern Hemisphere.
The genus contains three species found in California.
Jeannel (1963a: 54-58) revised the species. Soon after, Jeannel (1963b) published a supplement to his monograph and pointed out that his
A study of the type specimens of
This species is known from a few specimens collected in Tulare and Santa Clara Counties in California (Jeannel 1963a: 56).
This species is known only from the two original specimens collected at the type locality in south-central California.
This species is known from a few specimens collected near Springville in south-central California.
Western Hemisphere, with 44 species in eastern North America (42 species), most of them in the Appalachian region, and the state of Santa
Sokolov et al. (2004) reviewed the North American species and provided a key for the identification of all but four species (
This species is known only from the three original specimens collected at the type locality in central Texas.
This species is known only from Taney and Barry Counties in southwestern Missouri (Sokolov and Watrous 2008: 541).
This species is known from the original two specimens collected at the type locality in southeastern Kentucky (Sokolov et al. 2004: 208).
This species is known from a few specimens collected at the type locality on the Potomac River in Maryland and northern Virginia (Sokolov et al. 2004: 198).
This species is known from Monroe County (Sokolov et al. 2004: 209) and Polk County (Giachino 2011: 113) in southeastern Tennessee.
This species is known only from the type locality in southwestern North Carolina.
This species is known only from the type locality along the state line between Tennessee and North Carolina.
This species is known only from the holotype collected in western South Carolina near the Georgia border.
This species is known from the southeastern parts of Blount County in Tennessee and adjacent parts of Nantahala National Forest in Graham County, North Carolina (Sokolov and Carlton 2008: 42).
This species is known only from the holotype.
This species is known only from central and northeastern Blount County in Tennessee and adjacent parts of Swain County in North Carolina (Sokolov et al. 2007: 7).
This species is known only from two nearby localities in Gaston County, North Carolina, and York County, South Carolina (Sokolov et al. 2004: 210).
This species is known from two nearby localities in Burke County, western North Carolina (Sokolov et al. 2004: 211).
This species is known only from the holotype collected in central Texas.
This species is known only from the type locality in southwestern North Carolina (Sokolov et al. 2004: 212).
This species is known only from the holotype.
Jeannel (1963a: 76-77) reported that “Horn (1888: 27)” doubted the state provenance of the holotype and that the specimens in the USNM from Clayton, Georgia belong to
This species is known from several localities in Orange, Mecklenburg, and Cabarrus Counties in North Carolina (Sokolov et al. 2004: 200).
This species is known from southwestern Virginia and North Carolina, north of the French Broad River (Sokolov et al. 2004: 213).
This species is known only from the type locality in southwestern Alabama (Sokolov et al. 2004: 214).
This species is known from North Carolina, north of the French Broad River (Sokolov et al. 2004: 201), and presumably eastern Tennessee (Horn 1869a: 127). The records from the District of Columbia (Ulke 1902: 6), Virginia (Schwarz 1891: 24), South Carolina (Ciegler 2000: 55), and northeastern Georgia (Leng 1910: 73; Fattig 1949: 18) are probably in error.
This species is known only from the eastern part of Rich Mountain ridge, Blount County, eastern Tennessee (Sokolov and Carlton 2010: 12).
This species is known only from the original four specimens collected in Lawrence and Crawford Counties in southern Indiana (Sokolov et al. 2004: 202).
This species is known only from the type locality in southeastern Tennessee.
This species is known only from the type locality in northern Florida (Sokolov et al. 2004: 215).
This species is known only from the Great Smoky Mountains in Cocke, Monroe, and Sevier Counties, Tennessee (Sokolov et al. 2004: 216; Giachino 2011: 111).
This species is known only from Latimer County in eastern Oklahoma (Sokolov et al. 2004: 217).
This species is known only from the holotype collected in southern Tennessee.
This species is known in the Appalachians from western North Carolina, eastern Tennessee, and northwestern South Carolina (Sokolov et al. 2004: 219; Giachino 2011: 110).
This species is known only from Mount Magazine in western Arkansas (Sokolov et al. 2004: 221).
This species is known from two localities in Swain and Macon Counties in western North Carolina (Sokolov and Carlton 2010: 11).
This species is known only from high elevations in central Great Smoky Mountains National Park in eastern Tennessee and western North Carolina (Sokolov et al. 2004: 222).
This species is known only from Swain and Jackson Counties in North Carolina (Sokolov et al. 2004: 223).
This species is known only from the type locality in southwestern North Carolina.
This species is known from Avery and Wilkes Counties along the Blue Ridge Parkway in western North Carolina (Giachino 2011: 115).
This species is known from southwestern Blount County in Tennessee and Swain County in North Carolina in the Great Smoky Mountains National Park (Sokolov et al. 2007: 10).
This species is known only from the southern parts of the Ouachita National Forest in western Arkansas (Sokolov et al. 2004: 224).
This species is known only from the holotype collected in eastern Tennessee (Sokolov et al. 2004: 204).
This species is known only from the holotype collected in south-central Texas.
This species is known only from eight specimens collected at the type-locality cave.
This species is known from Tishomingo County in northeastern Mississippi, northern Alabama (Sokolov and Carlton 2010: 13; Giachino 2011: 111), Blount County in eastern Tennessee, Swain County in western North Carolina, and Dade County in northwestern Georgia (Sokolov et al. 2004: 205).
This species is known only from Latimer County in eastern Oklahoma (Sokolov et al. 2004: 225).
This species is known only from the type locality in central Arkansas (Sokolov et al. 2004: 226).
This species is known for sure only from the two original specimens collected in central Georgia. The records from York and Lancaster Counties in northern South Carolina (Ciegler 2000: 55) need confirmation.
This species is known only from the holotype collected in the central part of the Unicoi Mountains in North Carolina.
This true troglobite species is confined to caves in northeastern Alabama east of the Wills Creek anticline (Barr 1995: 243).
This species is known for sure only from the type locality in northern Virginia (Sokolov et al. 2004: 207). The record from “West Virginia” (Bousquet and Larochelle 1993: 157) needs confirmation.
Three species in eastern North America.
Sokolov and Carlton (2008: 40) provided a key for the identification of all species.
This species is known only from the holotype collected in northwestern South Carolina. The records from North Carolina and northeastern Georgia (Barr 1995: 245) refer to
This species is known only from the type locality in southwestern North Carolina.
This species is known from Giles and Botetourt Counties in Virginia (Sokolov and Carlton 2008: 40).
Two species in the Sierra Nevada of California.
Jeannel (1963a: 79) provided a key for the identification of the two species.
This species is known only from the holotype collected in eastern California.
This species is known only from the holotype collected in eastern California.
One troglobitic species in eastern North America.
One species in eastern United States.
Valentine (1932b) published a detailed description of the species enhanced by several accurate drawings.
This species is known only from the holotype collected in a cave three miles north of Lewisburg, near Maxwelton, in southeastern West Virginia. The cave has two rather small rooms connected by a narrow, descending, and tortuous passage. The specimen was found in the lower room, which was wet, muddy, and quite dark (Valentine 1932b: 1-2).
About 85 species (Lorenz 2005: 237-238) in the Nearctic (six species), Neotropical (five species), Australian (about 15 species), Palaearctic (about 50 species), and Afrotropical (five species) Regions arrayed in 12 genera:
Bousquet and Laplante (1997) revised the Western Hemisphere species and provided a key for their identification.
One species confined to the Pacific Coast of California and Baja California.
This species is found along the Pacific Coast from northern California to southern Baja California [see Bousquet and Laplante 1997: map 1].
Western Hemisphere, with four species in the Nearctic (four species, two of them endemic) and Neotropical (two species) Regions, including the West Indies.
This species is known from a few localities around Salton Sea in southern California, from the northern part of the Baja California Peninsula, and from the state of Sinaloa along the Pacific Coast in Mexico [see Bousquet and Laplante 1997: map 2].
This species is presently known from numerous specimens collected in two localities in Chaves County, New Mexico [see Bousquet and Laplante 1997: map 2].
This species ranges from south-central Oregon to south-central Kansas, south to western Texas near the Rio Grande, southeastern Arizona [see Bousquet and Laplante 1997: map 3], and southwestern California (LeConte 1874b: 44). One specimen without specific locality is known from Nebraska (Bousquet and Laplante 1997: 714).
This species is known from New Jersey, southern South Carolina (Ciegler 2000: 56), the Florida Peninsula including the Keys, some islands in the Greater Antilles, southeastern Texas, Chiapas in Mexico, southern New Mexico (Otero County, CMNH), and southeastern Arizona [see Bousquet and Laplante 1997: map 2]. The species is also found in Colombia (Bousquet and Laplante 1997: 710) and Venezuela (Chaudoir 1872a: 36).
About 55 species in two subgenera:
About 50 species in the Nearctic (one species), Australian (17 species), Palaearctic (about 30 species), and Afrotropical (four species) Regions.
This species is known only from a few localities in southwestern Louisiana and eastern Texas south to the Rio Grande [see Bousquet and Laplante 1997: map 1]; it was also collected at least twice at Atlantic City, New Jersey (G.E. Horn in Smith 1890: 79) in the xix Century.
About 220 species in the Nearctic, Oriental, and Palaearctic Regions arrayed in two tribes:
Northern Hemisphere, with about 215 species arrayed in four subtribes following Zamotajlov (2002):
Darlington (1938) and Lindroth (1961a) reviewed the North American species and provided keys for their identification. Subsequently to Lindroth’s work, one species (
Northern Hemisphere, with about 115 species in the Nearctic (four species) and Palaearctic (about 105 species) Regions arrayed in nine genera:
Twenty-three species (Lorenz 2005: 241-242) in the Nearctic (four species) and Palaearctic (19 species in Asia only) Regions arrayed in two subgenera:
Six species in the Nearctic (four species) and Palaearctic (
Marek and Kavanaugh (2005) revised the North American species and provided a key for their identification.
The range of this species extends from northern Northwest Territories and northern Yukon Territory south to central Oregon (Darlington 1938: 150), north-central Utah, and southern Colorado [see Marek and Kavanaugh 2005: Fig. 10].
This species ranges from western Montana (Russell 1968: 47) to Vancouver Island (Lindroth 1961a: 188), south to the northern part of the Coast Ranges in California and the Sierra Nevada (Darlington 1938: 149; Marek and Kavanaugh 2005: 160).
This species ranges from the Skeena River drainage in central British Columbia (Lindroth 1961a: 189) south to the Coast Ranges and the Sierra Nevada in central California (Darlington 1938: 151; Marek and Kavanaugh 2005: 164).
This eastern species is found from Cape Breton Island (Lindroth 1961a: 189) to the Saint Lawrence Valley in southern Quebec (Larochelle 1975: 81), south to northeastern West Virginia (Randolph County, CMNH). Fossil remnants from the Early Wisconsinan have been unearthed in southern Ontario (Morgan and Morgan 1981: 1107).
Northern Hemisphere, with about 80 species in the Nearctic (eight species) and Palaearctic (about 75 species, only six of them occurring in Europe) Regions arrayed in ten genera:
Sixteen species in the arctic, subarctic, boreal, and temperate areas of the Nearctic (seven species) and Palaearctic (13 species) Regions. Four species are Holarctic.
This western species ranges from southeastern Alberta to Vancouver Island, north to the Gulf Coast of Alaska, including the Aleutian Islands (Lindroth 1961a: 181), south to Mono County in the Sierra Nevada (CAS) and mountains in southwestern Colorado (Darlington 1938: 162-163). The record from Kamchatka (Eschscholtz 1823: 104) is probably in error since the species is not listed from the Palaearctic Region by Zamotajlov (2003b: 284-285).
Pohl (1998) reported that members of this species can be segregated into a “coastal” and “inland” morphs and that hybridization occurs between the coastal form of
This species is found from Newfoundland (Lindroth 1955a: 83) to the Okanagan River in south-central British Columbia, north to the Peace River in northeastern British Columbia (Lindroth 1961a: 181), south to eastern South Dakota (Kirk and Balsbaugh 1975: 21), northern Minnesota (Lindroth 1963a: Fig. 61), and the Pontiac region in southwestern Quebec (Lindroth 1961a: 181); isolated in high mountains in Colorado (Darlington 1938: 160).
This species ranges from Newfoundland (Lindroth 1955a: 83) to the Chukchi Sea coast in northwestern Alaska (Lindroth 1961a: 183), south to north-central Washington (Pohl 1998: 690), northern Minnesota (Clearwater and Lake Counties, CNC), the upper peninsula of Michigan (Escanaba, MCZ), and north-central Maine (Larochelle and Larivière 1990a: 28; Piscataquis County, CNC). The species is also known from the Far East (Zamotajlov 2003b: 285). Fossil remnants, dated between 10,400 and 15,400 years B.P., have been unearthed in Cape Breton Island in Nova Scotia (Miller 1997: 250) and central Iowa (Schwert 1992: 76); others from a Plio-Pleistocene sequence have been found in northwestern Greenland (Böcher 1995: 23).
This Holarctic species is found in easternmost Siberia (Zamotajlov 2003b: 285) and on this continent from Alaska, south of the Arctic Circle and including the Aleutian and Kodiak Islands (Lindroth 1961a: 186), to Newfoundland (Lindroth 1955a: 85), south to Mont Katahdin in Maine, the White Mountains in New Hampshire (Darlington 1938: 169), the Adirondack Mountains in northeastern New York (Notman 1928: 221, as
This widely distributed species ranges from Newfoundland (Lindroth 1955a: 82) to at least the Okanagan Valley in British Columbia, north to southern Northwest Territories (Lindroth 1961a: 180), south to southern Arizona (Darlington 1938: 158), the Sacramento Mountains in New Mexico (Fall and Cockerell 1907: 158), northern Oklahoma (Tulsa and Cimarron Counties, CNC), and northeastern Florida (Darlington 1938: 158). The record from “Texas” (Wickham 1896c: 132) needs confirmation.
The range of this species remains largely undocumented. It was reported by Zamotajlov (2003c: 242) from eastern Siberia, including the Kurils and Commander Islands, southern Alaska, including the Aleutian Islands, the upper peninsula of Michigan along Lake Superior (Casey 1920: 186, as
This circumpolar species ranges from Greenland (Böcher 1988: 8) west to Iceland (Zamotajlov 2003c: 241), south on this continent to northern Maine (Darlington 1938: 166; Dearborn and Donahue 1993: 5), northern New Hampshire (Coos County, Ross T. Bell pers. comm. 2008; Leng and Beutenmüller 1895: 75), northern Michigan (Lindroth 1961a: 184), northern Minnesota (Petrice et al. 2002: 9; Gandhi et al. 2005: 926), and southern Montana (Hatch 1933a: 7); isolated on high mountains in Colorado (Wickham 1902: 235; Zamotajlov 2003c: 241). Three specimens labeled “W[ashington]T[erritory]” are known (MCZ). Some of the records listed here may in fact refer to
According to Zamotajlov and Isaev (2006), this species is represented by four subspecies in the Palaearctic Region: the nominotypical subspecies,
One species in boreal and northern temperate regions of North America.
Bousquet and Grebennikov (1999) redescribed the species and illustrated the male and female genitalia.
With the exception of accessory setae on the pronotum, elytra, and mesosternum, this species is structurally very similar to those of
This species ranges from New Brunswick to Great Slave Lake in Northwest Territories, south to the foothills of the Rocky Mountains in southern Alberta, northeastern Minnesota (Gandhi et al. 2005: 926), northeastern Ohio (Ashtabula and Trumbull Counties, Harry J. Lee, Jr. pers. comm. 2008), northeastern Pennsylvania (Pike County, CMNH), and Connecticut (Krinsky and Oliver 2004: 396) [see Bousquet 1987a: map 2].
Five species placed in one genus.
Five species in Siberia (four species) and western North America (one species).
This rarely collected species is known from Yukon Territory (CNC), northeastern Alaska (Lindroth 1961a: 191), and the Kodiak Island (Dave H. Kavanaugh pers. comm. 2009) south to northwestern Oregon (Westcott et al. 2006: 8) and northwestern Montana (Edwards 1975: 50).
This subfamily includes a single tribe.
Six species arrayed in three genera:
Three species, two in tropical Africa and one in North America, Mexico, southern Europe, and the Middle East.
The species found in North America was treated in Lindroth’s (1961a: 175) monograph.
The range of this species, also known as the “stinking Beetle”, extends from southern Quebec, as far north as
One North American species.
The species was treated in Lindroth’s (1961a: 175-176) monograph.
This species is known from scattered localities from the Lac Saint-Jean area in Quebec (Larochelle 1975: 99) to Vancouver Island, north to the Skeena River in northwestern British Columbia (Lindroth 1961a: 176), south to the San Gabriel and San Bernardino Mountains in southern California (Noonan 1967: 92), southern Arizona (McCleve 1975: 176), southwestern New Mexico (Grant County, USNM), and the upper peninsula of Michigan along Lake Superior (LeConte 1846a: 154). The record from “Illinois” and “South Dakota” (Bousquet and Larochelle 1993: 158) need confirmation.
Worldwide, with about 735 species in 47 genera. These genera are placed in five tribes:
Three species in the genera
Two species in western North America.
Bousquet and Goulet (1990) commented on the structural differences between the two species and on the three subspecies of
This subspecies ranges from southwestern British Columbia (Lindroth 1961a: 8) to San Luis Obispo County in California along the Coast Ranges and, usually at low elevation, to Tulare County in the Sierra Nevada (Bousquet and Goulet 1990: 17, Fig. 6).
This subspecies is found in the Sierra Nevada near Lake Tahoe and in Yosemite National Park (Bousquet and Goulet 1990: 17, Fig. 6).
This subspecies occurs in the southern part of the Sierra Nevada from Kings Canyon and Sequoia National Park to northern Kern County (Bousquet and Goulet 1990: 17, Fig. 6).
This species is known from three localities in Idaho County in north-central Idaho (James C. Bergdahl pers. comm. 2008) and in Ravalli County in western Montana (Moore 2008: 81).
About 180 species arrayed in 21 genera in the Neotropical Region (about 120 species of which five extend into southwestern United States), Asia (about 40 species in five endemic genera, most species in the Oriental Region, a few in New Guinea, Japan, Taiwan, and Nepal), and Afrotropical Region (about 20 species in two endemic genera).
All North American species have been covered in the revision of the ozaenine species of southwestern United States and of selected ones from Mexico by Ball and McCleve (1990). The work includes a key for the identification of ten species.
About 50 species ranging from southwestern North America to Chile and Uruguay.
Recent molecular data analyses (Moore 2008) suggest that
This species ranges from southeastern California (San Bernardino County, CMNH) to southern Texas (Cameron County, CMNH), south to Brazil; it is also known from Cuba and from the Tres Marias Islands off the Pacific Coast of Mexico (Ball and McCleve 1990: 95).
About 40 species extending collectively from southwestern United States to Brazil and Paraguay.
This subspecies is known from Arizona and the states of Durango and Sonora in northwestern Mexico (Ball and McCleve 1990: 92).
This species is known from southern Arizona, western Durango, northern Sonora, and Baja California (Ball and McCleve 1990: 91).
Six species in the Neotropical Region, of which one extends into southern United States.
This species ranges from southeastern Texas (Ball and McCleve 1990: 87) to Guatemala (Bates 1881: 27).
Twelve Neotropical species, of which one extends into southwestern United States.
This species ranges from southern Arizona to northern Argentina, including the Cayman Islands in the West Indies (Ball and Shpeley 1990: 814-815).
About 650 species arrayed in two tribes:
About 535 species arrayed in nine genera. The genera are classified in four subtribes following Erwin (1970a: 27-38):
About 305 species arrayed in three genera:
About 300 species (Lorenz 2005: 17-20) in the Nearctic (50 species), Neotropical (about 50 species, several shared with North America), Australian (one species in New Guinea), Oriental, Palaearctic (about 120 species), and Afrotropical Regions. These species are arrayed in nine subgenera:
Erwin (1970a) revised the North and Middle American species. No new species from the region have been described subsequently.
Eighty-three species in the temperate, subtropical, and tropical regions of the Western Hemisphere and one species (
In a cladistic analysis based on adult characters carried out by Erwin (1970a: 170),
The records of
This species is found from northeastern Kansas south to southeastern New Mexico and San Luis Potosí in Mexico [see Erwin 1970a: Fig. 450].
This species extends from Connecticut and “Rhode Island” (Sikes 2003: 8) to southern Minnesota and northeastern Nebraska, south to the Rio Grande in south-central New Mexico and southeastern Texas, and the Florida Keys [see Erwin 1970a: Fig. 201].
This species ranges from Massachusetts to western Wyoming and western Colorado, south to central New Mexico, southeastern Texas, southeastern Mississippi (George County, Drew A. Hildebrandt pers. comm. 2008), and the Florida Keys [see Erwin 1970a: Fig. 200]. The populations in the northeast are isolated from the remaining ones but this is probably the result of inadequate collecting.
This species is found only in the southern parts of the United States from southern South Carolina to southern Florida, west to southern Arkansas and eastern Texas [see Erwin 1970a: Fig. 199]. The record from New Jersey (Smith 1910: 212) needs confirmation.
This species is known from east-central Texas, Latimer County in Oklahoma [see Erwin 1970a: Fig. 89], and from southwestern Mississippi (Adams County, Drew A. Hildebrandt pers. comm. 2008).
This species ranges from western New York to southeastern Minnesota, south to southeastern Texas, east-central Louisiana (West Feliciana Parish, Igor M. Sokolov pers. comm. 2009), and northern Florida [see Erwin 1970a: Fig. 88].
This species is known only from the holotype.
This species is known only from a few specimens collected in eastern Michigan, Missouri, northwestern Louisiana (Bossier Parish, Igor M. Sokolov pers. comm. 2009), and northwestern Mississippi [see Erwin 1970a: Fig. 90].
The range of this species extends from New Brunswick to east-central South Dakota (Kirk and Balsbaugh 1975: 40), south to northwestern Arkansas, northwestern Tennessee (Cheatham County, CMNH), and Virginia. The species is also known from a number of localities in the southern part of the Rocky Mountains, in Colorado and New Mexico [see Erwin 1970a: Fig. 391]. The records from Georgia (J.E. LeConte 1849: 25; Fattig 1949: 42), southwestern Alabama (Löding 1945: 23), and “Utah” (Horn 1872c: 384; Erwin 1970a: 147) need confirmation.
This species ranges from southwestern New Brunswick (Webster and Bousquet 2008: 23) to south-central British Columbia, south to southwestern Idaho, central Colorado, Missouri, and New Jersey along the Atlantic Coast [see Erwin 1970a: Fig. 392].
This species is known from western New York and southern New Hampshire (Hillsborough, Rockingham, and Strafford Counties, Ross T. Bell pers. comm. 2008) south to North Carolina, from several localities in southern Indiana [see Erwin 1970a: Fig. 414], and from Waukesha County in southeastern Wisconsin (Messer 2010: 34).
This species is known only from the original two specimens, one collected at the type locality and the other at an unspecified locality in Florida (Erwin 1970a: 150).
The range of this species extends from Maine (Larochelle and Larivière 1990a: 33) and southern Quebec to western South Dakota, south to central Texas and central Georgia (Butts County, CMNH; Horn and Ulyshen 2009: 121) [see Erwin 1970a: Fig. 417].
This species is known only from the original five specimens collected at the type locality in southwestern Alabama.
This species ranges from North Carolina to the Florida Keys, west to southern Alabama [see Erwin 1970a: Fig. 416]; also known from one locality in central Missouri (Erwin 1970a: 153).
This species is known from southeastern Michigan southeastwards to southern Florida, including the Keys, and southwestwards to east-central and southwestern Texas [see Erwin 1970a: Fig. 393]. The records from southeastern South Dakota (Kirk and Balsbaugh 1975: 39) and southwestern Ohio (Wright and Whitehouse 1941: 70, as
This species is known from a small area along the Atlantic Coast from New Hampshire to New Jersey [see Erwin 1970a: Fig. 415]. One specimen labeled from Crescent City, Florida is probably mislabeled (Erwin 1970a: 155).
This species ranges from northern California to eastern Utah, south to Guatemala (Erwin 1973b: 82) and northern Baja California [see Erwin 1970a: Fig. 197]. Specimens labeled from “Arkansas” and “Kansas” are known but could be mislabeled (Erwin 1970a: 87).
This species is known from the type locality and “Arizona” (Erwin 2011b: 285).
This species ranges from southeastern Arizona south to Guerrero, Mexico [see Erwin 1970a: Fig. 276].
This species ranges from northern Arizona and central New Mexico south to Durango City, Mexico [see Erwin 1970a: Fig. 219].
This species is confined to northern and central Florida [see Erwin 1970a: Fig. 302].
This species ranges from Cumberland County in Nova Scotia (Majka and Gilhen 2008: 1) to southern Manitoba, south to eastern Texas, northwestern Louisiana (Natchitoches Parish, Igor M. Sokolov pers. comm. 2009), southern Alabama, northwestern South Carolina (Ciegler 2000: 35), and Virginia along the east coast [see Erwin 1970a: Fig. 330]. The species is also recorded from an unspecified locality in New Mexico (Erwin 1970a: 128). The records from “Georgia” (J.E. LeConte 1849: 25, as
The range of this species extends from eastern Arizona to the Pacific Coast in southern California and Baja California Norte [see Erwin 1970a: Fig. 365].
This widely distributed species ranges from southwestern New Brunswick (Webster and Bousquet 2008: 23) to south-central British Columbia, south to southwestern California, the state of Tabasco in Mexico, and central Florida [see Erwin 1970a: Fig. 369].
This species ranges from northern California to southern Baja California Sur and southeastern Arizona [see Erwin 1970a: Fig. 332].
The range of this species extends from the Pacific Coast in southern California to northeastern Texas, north to northeastern Colorado, south to San Luis Potosí in central Mexico [see Erwin 1970a: Fig. 364].
This species is known only from Arizona [see Erwin 1970a: Fig. 278].
This species ranges from western Arizona to the Texas Panhandle and southeastern Texas [see Erwin 1970a: Fig. 254]; also known from northwestern Mississippi (Coahoma County, Drew A. Hildebrandt pers. comm. 2008).
This species is found west of the Appalachian Mountains from western New York to eastern Wyoming, including southwestern Wisconsin (Messer 2010: 34) and est-central Minnesota (Gandhi et al. 2011: 673), south to northeastern Mexico [see Erwin 1970a: Fig. 253].
This species ranges from the Atlantic Coast in New England to south-central British Columbia, south to southern California, northeastern Mexico, and central Florida [see Erwin 1970a: Fig. 331].
This species occurs from eastern Washington to northern Illinois, south to Guatemala and the southern part of the Baja California Peninsula [see Erwin 1970a: Fig. 252].
This species is restricted to the Coastal Plain where it ranges from North Carolina to central Florida, west to “Mississippi” (Drew A. Hildebrandt pers. comm. 2007) [see Erwin 1970a: Fig. 255].
This species ranges from southeastern Maine (Majka et al. 2011: 47) and southern Quebec to southwestern South Dakota, south to northern Texas, “Mississippi” (Drew A. Hildebrandt pers. comm. 2007) and, east of the Appalachian Mountains, to Long Island, New York [see Erwin 1970a: Fig. 303].
This species ranges from southwestern Maine (Majka et al. 2011: 48) to Wisconsin (Messer 2010: 34), including northeastern Ohio (Ashtabula County, Foster F. Purrington pers. comm. 2009), south to Maryland (Kent County, CMNH) [see Erwin 1970a: Fig. 301]. The records from “Georgia” (J.E. LeConte 1849: 25) and Missouri (Summers 1873: 133) are likely in error.
This species ranges from southeastern Virginia (Norfolk, Robert L. Davidson pers. comm. 2012) to eastern Oklahoma, south to southeastern Texas and southwestern Florida [see Erwin 1970a: Fig. 366]. The records from New York (LeConte 1846b: 203), New Jersey (Smith 1910: 212), southeastern Iowa (King 1914: 322; Hendrickson 1930: 94), and Missouri (Summers 1873: 133) need confirmation.
This species is restricted to the Great Plains and ranges from northeastern Iowa (Black Hawk County, Doug A. Veal pers. comm. 2009) to north-central Colorado, south to Chihuahua in northern Mexico and southeastern Texas [see Erwin 1970a: Fig. 279]. The specimens collected near Ithaca, New York (Erwin 1970a: 114), are possibly strays or mislabeled.
This species is known from a few specimens collected in the Gulf Plain in eastern Texas [see Erwin 1970a: Fig. 368].
The range of this species extends from Massachusetts to south-central British Columbia, south to southern California along the Pacific Coast, north-central Utah, central Kansas, southeastern Texas, southern Mississippi (George and Wilkinson Counties, Drew A. Hildebrandt pers. comm. 2008), and southern Florida [see Erwin 1970a: Fig. 251]. At least one specimen simply labeled from Arizona is known (Erwin 1970a: 101); the record from southwestern New Mexico (Fall and Cockerell 1907: 160, as
This species ranges from southern Quebec to southeastern Wyoming (Lavigne 1977: 44), south to southwestern Texas and northern Florida [see Erwin 1970a: Fig. 329].
This species is known so far only from the Central Valley in California between the Sierra Nevada and the Coast Ranges [see Erwin 1970a: Fig. 367].
This species ranges from southern Utah to the Rio Grande in southwestern Texas, south to the Mexican High Plateau in Jalisco [see Erwin 1970a: Fig. 220].
This species ranges from northeastern Washington to southern California [see Erwin 1970a: Fig. 218].
This species is mainly restricted to the Great Plains and ranges from southwestern Ohio to western Kansas south to southern Oklahoma and western Arkansas [see Erwin 1970a: Fig. 198].
This widely distributed species ranges from southeastern New York to southern Nebraska (Franklin County, Foster F. Purrington pers. comm. 2010), north to southwestern Wisconsin (Messer 2010: 34) and Michigan, south to Panama, southern Florida, and Cuba, west to the coast of the Gulf of California [see Erwin 1970a: Fig. 166].
This species ranges from southern Sonora in Mexico to Colombia [see Erwin 1970: Fig. 168]. It was also recorded from “Texas” (Erwin 2011b: 277).
This species extends from northern Arizona, southern Nevada, and southern California south to the southern part of the Baja California Peninsula and Chiapas in Mexico [see Erwin 1970a: Fig. 169]; also recorded from Nicaragua (Erwin 2011b: 292). The record from Cuba (Jacquelin du Val 1857: 12) must be in error.
This species extends along the Pacific Coast from central Oregon to the southern extremity of the Baja California Peninsula, and from western Arizona to southwestern Oklahoma (Kondratieff et al. 2005: 171), south to Oaxaca, Mexico [see Erwin 1970a: Fig. 109].
This species ranges from east-central Texas south at least to northern Colombia (Dejean 1831: 428) [see Erwin 1970a: Fig. 110].
This species ranges from Massachusetts to “Minnesota,” south to the Rio Grande River in southern Texas and west-central Florida [see Erwin 1970a: Fig. 108]. The species is quite common west of the Mississippi Basin and south of latitude 40°, but known only from a few isolated localities outside this area. The record from southeastern Alberta (Erwin 1970a: 62) is possibly based on a mislabeled specimen or a stray; that from Riverside County, California (Dajoz 2007: 19) is probably in error.
Worldwide, with about 19,600 species arrayed here in two supertribes for convenience:
Worldwide, with about 6,400 species arrayed in the following 24 tribes:
About 85 species (Lorenz 2005: 247-248) in the Nearctic (two species, none endemic), Neotropical (11 species), Australian (16 species), Oriental (11 species), Palaearctic (six species), and Afrotropical (40 species) Regions. These species are arrayed in nine genera:
About 40 species (Lorenz 2005: 247) in temperate, subtropical, and tropical areas of the Nearctic (two species), Neotropical (9 species), Australian (15 species), Oriental (eight species), Palaearctic (four species), and Afrotropical (three species) Regions.
Allen (1969) reviewed the Western Hemisphere species.
This species is known from southern Arizona, the Baja California Peninsula (Allen 1969: 146), and the state of Sonora (CAS).
This species ranges from east-central South Dakota to central New York, south to southern Florida and southeastern Texas (Allen 1969: 147). The record from Cuba (Jacquelin du Val 1857: 18) needs confirmation.
Worldwide, with approximately 810 species in about 45 genera. The tribe is underrepresented in the Northern Hemisphere with around 75 species (less than 10% of the world fauna). Only five species, all in the genus
Members of this tribe are often included in the tribe
About 210 species in the Nearctic (44 species), Neotropical (about 150 species), and Australian (about 20 species) Regions.
Allen (1972) revised the North American species though he left out more than 20 species because their type series were not located or included only females. These species were studied by Bousquet (2006a) leading to several new synonymies.
This species is known only from the type locality in southeastern Texas.
This species has been recorded from southeastern South Carolina (Ciegler 2003: [2]), northern Georgia (J.E. LeConte 1849: 26; Fattig 1949: 28), all over Florida except the Keys (Peck and Thomas 1998: 19), southwestern Alabama (Löding 1945: 17), Mississippi (Snodgrass and Cross 1983: 18), one locality (probably Mittie in Cherokee County) in eastern Texas (see Allen 1972: 94), and the Bahamas (Peck and Thomas 1998: 19). The record from “Arkansas” (Wickham 1896b: 43) needs confirmation; those from the District of Columbia (Ulke 1902: 7) and southwestern Ohio (Wright and Whitehouse 1941: 70) are probably in error.
This species is known from a few specimens collected in Maryland (Allen 1972: 110; CNC), eastern Alabama, and southeastern Oklahoma (Bousquet 2006a: 151-152).
This species is known from several localities in northern and central Florida and one locality in central Arkansas [see Allen 1972: Fig. 176]. The records from South Carolina (Kirk 1969: 11) and Georgia (Fattig 1949: 28, also as
This species is known from Dubois and Vigo Counties in western Indiana, from an unspecified locality in Arkansas (Allen 1972: 84), and from Noxubee County in eastern Mississippi (CMNH).
This species is known from the Bahamas and along the Coastal Plain from southern Florida west to eastern Texas, north to northeastern Louisiana [see Allen 1972: Fig. 171]. The record from southern Georgia (Fattig 1949: 27, as
This species is known only from the original 13 specimens collected at the type locality.
This species is known from northeastern Arkansas south to southeastern Louisiana, east to eastern Mississippi (Noxubee County, CMNH) and west to east-central Texas [see Allen 1972: Fig. 170]; also recorded from South Carolina (Kirk 1969: 11; Ciegler 2000: 59). One specimen simply labeled from Kentucky is known (Allen 1972: 98)
This species is known from a few localities in central Florida and southern Alabama [see Allen 1972: Fig. 174, as
This species is known only from the holotype (Bousquet 2006a: 149).
This species is known only from the type series.
This species is known from southeastern Georgia to central Florida including the Panhandle (Peck and Thomas 1998: 19) [see Allen 1972: Fig. 175, as
This species is known from a few localities in central Georgia, northern Florida (Peck and Thomas 1998: 19), northeastern Alabama (Allen 1972: 108), and northeastern Arkansas [see Allen 1972: Fig. 174].
This species is known only from the holotype collected in Georgia from an unspecified locality.
This species is known from two unspecified localities in Arkansas and Louisiana (Allen 1972: 102).
This species is known along the Coastal Plain from North Carolina and South Carolina, west to southern Louisiana and eastern Arkansas [see Allen 1972: Fig. 175]. Specimens simply labeled from Georgia are known (Allen 1972: 87). The records from Missouri (Summers 1873: 134), central Kansas (Knaus 1885: 58), and southeastern Texas (Casey 1918: 384) need confirmation.
This species is found along the Coastal Plain from northeastern Florida to central Louisiana and east-central Arkansas [see Allen 1972: Fig. 172]. At least one specimen simply labeled from Texas is known (Allen 1972: 78).
This species is known for sure only from the four original specimens. The records from “Pennsylvania” (LeConte 1853a: 252), South Carolina (Kirk 1969: 11), Georgia (J.E. LeConte 1849: 26; Leng 1910: 73; Fattig 1949: 28), and southwestern Alabama (Löding 1945: 17) need confirmation.
[AL, GA, PA, SC]
Dejean’s name was listed as synonym of
This species ranges from Virginia (Virginia Beach County, VMNH) to southern Florida, west to southeastern Texas (Casey 1918: 389, as
This species inhabits the Coastal Plain from southeastern Virginia (Norfolk, Robert L. Davidson pers. comm. 2008) to central Florida, west to southeastern Texas (Brazoria County, Brian Raber pers. comm. 2010) [see Allen 1972: Fig. 162], including western Arkansas (Polk and Garland Counties, Robert L. Davidson pers. comm. 2012); also recorded from Cuba (Darlington 1934: 91).
This species is at present known only from the holotype which Chaudoir (1868b: 344) believed had been collected in Texas.
This species is found from southeastern Pennsylvania to Iowa (Wickham 1888: 82; King 1914: 323; Jaques and Redlinger 1946: 297), south to southern Louisiana and central Florida including the Panhandle (Peck and Thomas 1998: 19) [see Allen 1972: Fig. 160]. Old specimens simply labeled from Massachusetts and Oklahoma are known (Allen 1972: 144).
This species is known from southwestern Ohio and northeastern Kentucky to western Illinois, south to north-central Texas, southern Alabama, and the Florida Panhandle (Peck and Thomas 1998: 19) [see Allen 1972: Fig. 168]. One specimen simply labeled from “South Carolina” is known (Allen 1972: 156).
This species is known from a few localities in Maryland, the District of Columbia, northern Ohio, Mississippi (Drew A. Hildebrandt pers. comm. 2007), and Alabama [see Allen 1972: Fig. 159]. The record from “Illinois” (Bousquet and Larochelle 1993: 161) needs confirmation.
This species ranges along the Coastal Plain from Rhode Island and Connecticut (William L. Krinsky pers. comm. 2008) to central Florida, west to southern Louisiana and north along the Mississippi River drainage to east-central Illinois [see Allen 1972: Fig. 155]. The record from southwestern Ohio (Dury 1902: 113) needs confirmation.
This species is known along the Coastal Plain from southern North Carolina to central Florida, west to northeastern Louisiana and southern Arkansas, north along the Mississippi River drainage to southwestern Indiana and southwestern Illinois (Allen 1972: 147-148, Fig. 156).
This species ranges from southern Ontario (Lindroth 1966: 540) south to southeastern Louisiana (Chaudoir 1843b: 766), west to eastern Kansas [see Allen 1972: Fig. 167]. The records from New Jersey (Smith 1910: 206), District of Columbia (Ulke 1902: 7), South Carolina (Kirk 1969: 11; Ciegler 2000: 59), “Georgia” (J.E. LeConte 1849: 26), Florida (Frost 1964: 138), and southwestern Alabama (Löding 1945: 17) need confirmation.
This species ranges from northern Indiana to eastern Iowa, south to northern Louisiana and southwestern Alabama [see Allen 1972: Fig. 161). One specimen simply labeled from Florida is known (Allen 1972: 153).
This species is known from one locality in southwestern Indiana, several localities in Arkansas, and one locality in Noxubee County in eastern Mississippi (CMNH) [see Allen 1972: Fig. 169].
This species is known from a few localities in northwestern Ohio, South Carolina, and southwestern Alabama [see Allen 1972: Fig. 165]. Specimens simply labeled from Virginia and Arkansas are known (Allen 1972: 132).
This species ranges from Maryland (Erwin 1981b: 160) to southeastern Minnesota (Allen 1972: 137), including north-central Ohio (Lee 1994: 61) and southern Ontario (Lindroth 1966: 539), south to southeastern Louisiana and southern Georgia (Torres and Ruberson 2006: 32) [see Allen 1972: Fig. 158]. The record from “Connecticut” (Bousquet and Larochelle 1993: ) was based on a misidentified specimen of
This species is found from northern Sonora and southeastern Texas south to Brazil (Allen 1972: 34); also known from Haiti [see Allen 1972: Fig. 144].
This species ranges from southeastern Arizona (Allen and Ball 1980: 500) south to Morelos in Mexico [see Allen 1972: Fig. 154].
This species ranges from southern Texas (Allen and Ball 1980: 501) and central Sinaloa south to Panama [see Allen 1972: Fig. 153].
The range of this species extends from central New York (Bousquet 2006a: 152) to east-central Iowa (Johnson County, CMNH), including southern Ontario (CNC), south to southeastern Texas (Cameron County, CMNH), central Louisiana, and northeastern Alabama (DeKalb County, CMNH) [see Allen 1972: Fig. 166].
This species is known from three localities in southwestern Tennessee, eastern Arkansas (Allen 1972: 114, Fig. 165), and southern Louisiana (Chaudoir 1868b: 342).
This species is known only from George and Noxubee Counties in eastern Mississippi (Will and Liebherr 1998: 231).
This species ranges from Connecticut (Krinsky and Oliver 2001: 102) and southeastern Pennsylvania to southwestern Illinois, south to southern Louisiana, southwestern Alabama, and central South Carolina (Ciegler 2000: 60) [see Allen 1972: Fig. 167]. One specimen simply labeled from Florida is known (Allen 1972: 116).
This species is known from central and southern Florida (Peck and Thomas 1998: 19), the Bahamas (Turnbow and Thomas 2008: 13), Grand Cayman Island, and southern Mexico [see Allen 1972: Fig. 146]. One specimen simply labeled from Texas is known (Allen 1972: 42).
This species is known from a few localities in northwestern Louisiana (Casey 1918: 383, as
This species is found west of the Appalachians from central Kentucky south to eastern Texas and southern Alabama (Allen 1972: 130, Fig. 163). The record from northern Georgia (Fattig 1949: 27) is probably in error.
This species is found along the Coastal Plain from southern North Carolina to the Rio Grande in southeastern Texas [see Allen 1972: Fig. 159].
This species ranges from the District of Columbia to southern Illinois, north to northern Ohio (Lee 1994: 61), south to southeastern Texas and northern Florida including the Panhandle (Peck and Thomas 1998: 19) [see Allen 1972: Fig. 157]. The record from southeastern Pennsylvania (Rathvon 1869: 525) needs confirmation.
This species is found from North Carolina to northwestern Arkansas, including south-central Kentucky (Metcalfe County, Peter W. Messer pers. comm. 2012), south to eastern Texas and southern Florida [see Allen 1972: Fig. 164].
Twenty species in the Neotropical Region, one of them extending into southern United States.
Will (2005) provided a key for the identification of the species of this genus.
This genus-group taxon has been ranked as a subgenus of
This species ranges from southern Arizona and southern Texas along the Gulf of Mexico coast south to southern Mexico (Allen and Ball 1980: 532). The record from southwestern Ohio (Dury 1906: 257) must be in error or based on a stray.
Worldwide, with about 2,620 species. The North American fauna is represented by about 255 species (9.7% of the world fauna), of which six are adventive, arrayed in 38 genus-group taxa including the eastern endemic
Twenty-seven species in the Neotropical Region, one of them reaching southwestern United States. The species are arrayed in two subgenera:
Will (2002b) published a revision of all known species.
A cladistic analysis placed the genus
Fourteen species in the Neotropical Region, one of them reaching southwestern United States.
This species is known from southern Arizona, possibly also southeastern California as suggested by the type locality, Sonora in northwestern Mexico, and the southern part of the Baja California Peninsula [see Will 2002b: Fig. 17].
Seven species in the Neotropical Region, one of them reaching southwestern United States.
There is no taxonomic revision or available key for the identification of the species.
This species is known from southern Arizona (Schaeffer 1910: 393, as
About 140 species (Lorenz 2005: 266-269) in the Nearctic (13 species) and Palaearctic (128 species) Regions; five of the North American species extend into northern Mexico and Cuba. The species of
No review or identification key is available for the North American species and a revision of the genus is needed.
About 70 species (Lorenz 2005: 266-268) in the Nearctic (12 species, of which five extend into northern Mexico or Cuba) and Palaearctic (about 60 species) Regions.
Chaudoir (1876c) reviewed the species but his work is outdated. Lindroth’s (1966) monograph covers seven of the 12 North American species.
This species ranges from southwestern New Brunswick (Majka et al. 2007: 9) to the Black Hills in western South Dakota (Kirk and Balsbaugh 1975: 21), south to northern Chihuahua (Bates 1891a: 249), southern Texas (Johnson 1978: 67), and the Florida Panhandle (Okaloosa County, CNC); also recorded from Cuba (Darlington 1934: 91). The records from Arizona (Snow 1906b: 162, as
This species ranges from “Colorado” (LeConte 1858a: 28) to west-central Nevada (Bechtel et al. 1983: 474), south to the Baja California Peninsula (Horn 1894: 309) and “New Mexico” (Schaupp 1882c: 41). The record from est-central South Dakota (Kirk and Balsbaugh 1975: 21) needs confirmation.
The range of this species extends from southern Manitoba to the Fraser River in south-central British Columbia (Lindroth 1966: 481), south to southern California (Fall 1901a: 44) and the Mexico City environs on the Mexican Plateau (Ball and Shpeley 1992a: 52).
This species is known for sure only from the syntype. The records from northeastern Kansas (Horn 1872c: 385; Popenoe 1878: 78) are suspect.
Adults of this species are structurally similar to, and possibly conspecific with, those of
This species is found from western Ontario (CNC) to southeastern Alberta (Lindroth 1966: 484), south to southeastern Oregon (Harney County, James R. LaBonte pers. comm. 2009; Hatch 1953: 115), central Wyoming (Lavigne 1977: 47), western South Dakota (Kirk and Balsbaugh 1975: 21), and western Minnesota (Tinerella and Rider 2001: 320; Gandhi et al. 2005: 926). The record from northern British Columbia (Jarrett and Scudder 2001: 382) needs confirmation.
This species is known from western Oregon (Hatch 1953: 115, as
This species has been known for a long time under the name
This species ranges from southeastern Washington (Hatch 1953: 115) to southern California (Fall 1901a: 44; Moore 1937: 9) and southern Arizona (Wickham 1898: 300). The records from New Mexico (Wickham 1896a: 157; Fall and Cockerell 1907: 158) and northeastern Kansas (Popenoe 1878: 78) are probably based on misidentified specimens.
This widely distributed species ranges from Cape Breton Island (Lindroth 1954c: 303) to Vancouver Island, north at least to Fort Smith in southern Northwest Territories (Lindroth 1966: 483), south to “Oregon” (Hatch 1953: 116), northern New Mexico (Fall and Cockerell 1907: 158), the Texas Panhandle (Michels et al. 2010: 743), southwestern Oklahoma (Kondratieff et al. 2005: 173), northern Georgia (Fattig 1949: 26), and central South Carolina (Ciegler 2000: 61).
The lectotypes of
This species is known from southern California (Fall 1901a: 45).
This species is known yet only from north-central (LeConte 1853a: 254) and southeastern Colorado (Michels et al. 2008), “Kansas” (Horn 1872c: 385), and a few localities in central Mexico (Bates 1882a: 84).
This species ranges from southeastern Arizona (Casey 1913: 137, as
This species is known from southeastern Arizona (Snow 1906b: 162) and southern Texas (Snow 1906a: 141; Duval County, CMNH). The records from Kansas (Knaus 1901: 110; Snow 1903: 193) need confirmation.
Twenty-three species (Lorenz 2005: 269), mostly in the arctic and subarctic areas, in North America (one species), Asia (21 species), and eastern Europe (three species).
No revision or identification key is available for the species. Lindroth (1966: 476-478) covered the North American species.
This species ranges from the Anderson River delta in northern Northwest Territories (Lindroth 1966: 478) to the Titaluk River (North Slope County) in northern Alaska (CNC). The species is also known from a few localities in eastern Siberia (Alfimov and Berman 2009: Fig. 1). Fossil remnants of this species, from the early Pleistocene and Late Wisconsin, have been found in the lower Kolymian Basin in northeastern Siberia (Matthews 1974a: 208) and western Alaska (Matthews 1974b: 1365) respectively.
Six species in the temperate regions of eastern North America.
Will (1999) reviewed the species and provided a key for their identification.
This species is known from a few localities along the Atlantic Coastal Plain and Piedmont Plateau from central New Jersey (Smith 1910: 205) to central South Carolina (Ceigler 2000: 62). The record from “Pennsylvania” (LeConte 1873a: 316) needs confirmation.
This species ranges from southwestern Indiana (Blatchley 1910: 97; Schrock 1985: 354) to southeastern Iowa (Wickham 1911b: 6; Henry County, CNC), south to eastern Texas and central Florida (Will 1999: 272, Fig. 35). The record from North Carolina (Brimley 1938: 121) needs confirmation; that from “New York” (Wickham 1895a: 187) is likely in error.
This species ranges from southern Quebec (Larochelle 1975: 105) to southeastern Minnesota (Gandhi et al. 2005: 926), south to Missouri (Summers 1873: 145) and West Virginia (Will 1999: 270). The records from North Carolina (Brimley 1938: 121), South Carolina (Kirk 1970: 11), and Georgia (J.E. LeConte 1849: 26; Fattig 1949: 26) need confirmation.
This species ranges from Long Island in New York (Schaupp 1883b: 31) and New Jersey to central Missouri, north to northeastern Illinois, south to southeastern Louisiana (Saint Tammany Parish, Igor M. Sokolov pers. comm. 2009) and central Florida [see Will 1999: Fig. 34].
As far as known, this species ranges along the Coastal Plain from the coast of South Carolina to central Florida, west to east-central Texas, north along the Mississippi Valley to southern Indiana (Will 1999: 271). The records from “Iowa” (Jaques and Redlinger 1946: 297), “Illinois” (Wolcott 1900: 469), “New York” (Wickham 1895a: 187; Casey 1913: 144), New Jersey (Smith 1910: 205), and southeastern Pennsylvania (Rathvon 1869: 526, as
This species is known from a few localities along or near the Atlantic Coast from New Hampshire to Virginia (Will 1999: 268).
One species in the temperate regions of eastern North America.
This species is found from southern Ohio (Purrington and Stanton 1996: 43; Silverman et al. 2008: 733) south to “Texas” (LeConte 1858a: 28), southern Mississippi (Hancock County, CNC), and central Florida (Peck and Thomas 1998: 19). Bates (1882a: 83) recorded it from “Mexico.”
Three species in the temperate regions of eastern North America.
Darlington (1932) provided a key for the identification of the species.
This species is found along the Appalachian Mountains from southern West Virginia (Hoffman 1998: 36) to northern Georgia (Leng 1910: 73; Fattig 1949: 21) and northwestern South Carolina (Ciegler 2000: 63).
This species ranges from the Nova Scotia Peninsula (Schaupp 1882c: 41; Chritopher G. Majka pers. comm. 2008) to western Minnesota (Gandhi et al. 2005: 926), south along the Appalachian Mountains to northern Alabama (Löding 1945: 15), northern Georgia (Leng 1910: 73; Fattig 1949: 21), and northwestern South Carolina (Ciegler 2000: 63).
This species is known from the Great Smokies along the Tennessee and North Carolina border (Darlington 1932: 155; Barr 1969: 81) and Rabun Bald in northeastern Georgia (Fattig 1949: 22).
About 40 species in the temperate areas of the Nearctic (two species, one of them adventive) and Palaearctic Regions, arrayed in two subgenera, both represented in North America.
One North American species along the Pacific Coast.
Hacker (1968) redescribed the species.
This rarely collected species is known from one specimen collected along the west coast of Vancouver Island (Bousquet 2000: 81) and a few others collected along or near the coast of Oregon (Hacker 1968: 42).
Thirty-six species in Asia (29 species) and Europe (seven species), one of them adventive in eastern North America.
There is no taxonomic revision of the subgenus. The sole species in North America was included in Lindroth’s (1966: 443) monograph.
This European species is adventive in North America where it is known from a few localities in Cape Breton Island (Bousquet 1987a: 124), Prince Edward Island (CNC), and New Brunswick (Kent County, Christopher G. Majka pers. comm. 2011). The first inventoried specimen collected on this continent was found in Cape Breton Highland National Park in 1984 (Bousquet 1987a: 124). The previous North American record from Hemmingford, Quebec (Darlington 1940b: 252) was based on a mislabeled specimen (see Bousquet 1987a: 124).
Two Holarctic species in the arctic, subarctic, and boreal regions.
Lindroth (1966) redescribed both species and provided a mean for their identification through his key to Canadian species of
This Holarctic species ranges in the Palaearctic Region from northern European Russia to the Far East, south to Manchuria (Bousquet 2003d: 517), and in the Nearctic Region from Alaska (Lindroth 1966: 533) to Newfoundland (Lindroth 1955a: 88), south to high mountains in New England (Lindroth 1966: 533), the Black Hills in southwestern South Dakota (Robert L. Davidson pers. comm. 2008), and Wyoming (Lindroth 1966: 533). The record from “New York” (Notman 1928: 226) needs confirmation; that from Ohio (Everly 1927: 155) is in error. Fossil remnants of this species, dated between about 10,400 and 18,100 years B.P., have been unearthed in southeastern Iowa (Baker et al. 1986: 96) and Cape Breton Island in Nova Scotia (Miller 1997: 250); others, believed to be 2.0-2.5 million years old, have been found in Greenland (Bennike and Böcher 1990: 336; Böcher 1995: 28).
This Holarctic species ranges from the northern part of European Russia (Bousquet 2003d: 517) to the Arctic Slope in northern Alaska. The record from “Yukon Territory” (Ball and Currie 1997: 452) could not be confirmed.
Thirty-two species (Lorenz 2005: 265-266) in the temperate regions of North America (two species), Asia (29 species), and Europe (one species), arrayed in two subgenera (Bousquet 1999: 90):
Thirty-one species (Lorenz 2005: 266) in eastern North America (two species) and eastern Asia (29 species).
Allen (1980) revised the North American under the generic name
This species ranges from southwestern Connecticut (Krinsky and Oliver 2001: 104) to northeastern Illinois, north to the northwestern region of the lower peninsula of Michigan (Allen 1980: 20; Hatch 1925: 549), south to northern Mississippi (Union County, Drew A. Hildebrandt pers. comm. 2010), northern Alabama (Löding 1945: 15), and northwestern Georgia (Fattig 1949: 21). The records from southern Wisconsin (Rauterberg 1885: 16) and “Massachusetts” (Bousquet and Larochelle 1993: 168) need confirmation.
This species is found from “Nova Scotia” (Larochelle and Larivière 1990a: 29) to northwestern Minnesota (Allen 1980: 24), south to northern Alabama (Löding 1945: 15) and Georgia (Fattig 1949: 21, as
About 1,055 species (Lorenz 2005: 269-287, 289, as
Based on phylogenetic analyses from two nuclear gene sequences (wingless and 28SrDNA), Sasakawa and Kubota (2007) concluded that the genus
Thirteen species in the boreal and temperate areas of the Nearctic (three species, one of them adventive) and Palaearctic (11 species) Regions.
Bousquet and Webster (2004) reviewed the Nearctic species.
This subgenus was known until the late 1980s under the name
This species ranges from Cape Breton Island and Prince Edward Island to south-central British Columbia, south to the Lake Tahoe region in the Sierra Nevada, southern Colorado along the Rocky Mountains, northeastern Kansas (Popenoe 1877: 23, as
This species ranges from Nova Scotia (Christopher G. Majka pers. comm. 2004) to the Missouri River in southeastern South Dakota and eastern Nebraska, south to southwestern Texas (Dajoz 2007: 23, as
This European species is adventive in North America where it is known from southern Quebec, south of the Saint Lawrence River, eastern Ontario (near Avonmore, CNC), and northern Vermont (Bousquet and Webster 2004: 658). The first inventoried specimen collected on this continent was found in 1997 (Byers et al. 2001: 84). The previous record from Quebec (Lindroth 1966: 503) was based on a mislabeled specimen.
Twenty-eight species (Lorenz 2005: 275) in the boreal and temperate areas of the Nearctic (four species, one of them adventive) and Palaearctic (25 species) Regions.
Lindroth (1966) reviewed the Nearctic species.
This small
The range of this species extends from the Saguenay River in Quebec (Larochelle 1975: 101) to east-central British Columbia (Jarrett and Scudder 2001: 383), south to northeastern New Mexico (Fall and Cockerell 1907: 158; San Miguel County, MCZ), the Texas Panhandle (Robert L. Davidson pers. comm. 2008), and the District of Columbia (Ulke 1902: 6). The records from “Georgia” (J.E. LeConte 1849: 26) and southern Louisiana (Summers 1874a: 80) need confirmation.
This species occurs from Newfoundland (Lindroth 1955a: 91) to the Gulf Coast of Alaska (Lindroth 1966: 506), south at least to northeastern Washington (Pend Oreille County, CNC), eastern Kansas (Popenoe 1877: 23; Lindroth 1966: 505), and southwestern Virginia (Hoffman 1998: 39).
This European species is adventive in North America on both coasts where it is found in southeastern Newfoundland (Lindroth 1955a: 91; Larson and Langor 1982: 593) and the Vancouver area in British Columbia (Spence and Spence 1988: 158, Fig. 8). The first inventoried specimen collected on this continent was found in Newfoundland in 1937 (see Lindroth 1955a: 91).
Eighteen species (Lorenz 2005: 273) in the arctic (marginal), subarctic, boreal, and temperate areas of the Nearctic (six species, one of them circumboreal) and Palaearctic (12 species, including two in the Himalayas) Regions, with one species (
Tschitschérine (1900d) published a key to the species but his work is outdated. Lindroth (1966: 484-491) reviewed the North American species under his
This circumpolar species ranges from Iceland to the coast of Bering Sea (Bousquet 2003d: 487) and from Alaska, including the Aleutian, Kodiak, and possibly Pribilof Islands (Lindroth 1966: 487), to Newfoundland (Lindroth 1955a: 94), south to north-central Pennsylvania (LeConte 1867a: 346, as
This species ranges from southern British Columbia (Lindroth 1966: 489) south to northern Utah (Cache, Davis and Utah Counties, Robert L. Davidson pers. comm. 2012) and eastern Oregon (Hatch 1953: 118).
The range of this species extends from Newfoundland (Lindroth 1955a: 93) to central British Columbia, north to Fort Smith in southern Northwest Territories (Lindroth 1966: 487), south to the Black Hills in western South Dakota (Kirk and Balsbaugh 1975: 22), southeastern Iowa (Wickham 1911b: 6), and west-central Virginia (Hoffman 1998: 39).
This species ranges from Vancouver Island (Lindroth 1966: 491) and northern Idaho (Idaho County, CNC) south to central New Mexico (Bernalillo County, CMNH), southern Arizona (Wickham 1898: 300; Snow 1906b: 162), and southern California (Fall 1901a: 45; Moore 1937: 9).
This species ranges from western Newfoundland (Lindroth 1955a: 92) to “Montana” (Lindroth 1966: 490), south to northeastern New Mexico (San Miguel County, CNC), northeastern Kansas (Popenoe 1877: 23), and northeastern Georgia (Leng 1910: 73; Fattig 1949: 27); isolated in southwestern British Columbia, including Vancouver Island (Lindroth 1966: 490). The record from “Idaho” (Bousquet and Larochelle 1993: 170) needs confirmation. Lindroth (1966: 490) postulated that the best explanation for the presence of this species in southwestern British Columbia is that it was accidentally introduced into the area.
This species ranges from southeastern South Dakota (Kirk and Balsbaugh 1975: 22) to southwestern New York (Smith 1910: 205), south to Virginia (Nicolay and Weiss 1934: 211; Carrington 2002: 108), central West Virginia (Carrington 2002: 108), and central Missouri (Boone County, CMNH).
Eight species in the boreal and temperate regions of the Nearctic (three eastern species) and Palaearctic (five species, of which three occur in the Far East) Regions.
Lindroth (1966) reviewed the North American species. One new species was described subsequently by Goulet and Bousquet (1983).
The range of this species extends from Nova Scotia (Majka et al. 2007: 9) to south-central British Columbia, as far north as northeastern British Columbia (Lindroth 1966: 499-500) and southern Northwest Territories (CNC), south to eastern Washington (Hatch 1953: 117), northern Colorado (Haubold 1951: 704; Armin 1963: 224), southern Iowa (Union County, Foster F. Purrington pers. comm. 2011), and northeastern South Carolina (Ciegler 2000: 66). The record from southwestern California (Moore 1937: 9) is probably in error.
This species is restricted to a small area from New Brunswick (Webster and DeMerchant 2012: 6) to western Ontario, south to northern Wisconsin (Messer 2010: 37), northern Michigan (Dunn 1985a: 12), and northeastern Pennsylvania (Pike County, CMNH) [see Goulet and Bousquet 1983: map 1].
This species ranges from “Rhode Island” (Sikes 2003: 7) to “Minnesota” (Gandhi et al. 2005: 926), including southernmost Ontario (Lindroth 1966: 499, possibly only as strays), south to “Texas” (Schaupp 1882c: 42; Nicolay and Weiss 1934: 209), southeastern Louisiana (Lindroth 1966: 498), and southern Florida (Peck and Thomas 1998: 20).
Eighteen species (Lorenz 2005: 274-275) in the boreal and temperate regions of the Nearctic (two species) and Palaearctic (16 species) Regions.
Bousquet and Pilon (1984) commented on the structural differences between the two North American species.
This species ranges from Newfoundland to Vancouver Island, south to west-central Washington, central Colorado (Jefferson County, Robert L. Davidson pers. comm. 2008), east-central South Dakota, northern Illinois, and eastern Virginia [see Bousquet and Pilon 1984: Fig. 8].
This species ranges from eastern Newfoundland to northeastern Alberta, south to northern Colorado, northern Illinois, and southwestern North Carolina (Macon and Jackson Counties, CNC) [see Bousquet and Pilon 1984: Fig. 7].
Two species in the Appalachian region of eastern North America.
Bousquet (2006b) commented on the structural differences between the two species.
This species is known from northeastern Kentucky, eastern and central West Virginia, and southwestern Virginia (Bousquet 2006b: 159).
This species is found at high elevation in the Appalachian Mountains in North Carolina, Tennessee (Barr 1969: 72), Kentucky (Jackson County, Foster F. Purrington pers. comm. 2009), and northern Georgia (Fattig 1949: 24), including the Black, Bald, Great Balsam, and Great Smoky Mountains. The record from “Virginia” (Bousquet and Larochelle 1993: 172) needs confirmation.
One species in the Appalachian Mountains of eastern North America.
This species is endemic to the Appalachian Mountains from southwestern Virginia (Hoffman 1998: 38) to northern Georgia (Fattig 1949: 26) and northwestern South Carolina (Ciegler 2000: 67), including the Great Balsam and Great Smoky Mountains (Barr 1969: 80).
Nine species in the Pacific Northwest.
Bousquet (1985a) wrote a key for the identification of the species. Three new species were described subsequently (Bousquet 1992b; Bergdahl and Kavanaugh 2011).
This species is known from several localities in western Oregon. Two specimens simply labeled “Was” are also known (Bousquet 1985a: 258).
This species is known from western Washington (Bousquet 1985a: 259). The record from northwestern Oregon (Hatch 1953: 114) probably refers to
This species is known from the type locality in southern Washington and one locality in northwestern Oregon (Westcott et al. 2006: 8).
This species is known from Latah and Idaho Counties (CNC) in northern Idaho and Mineral County in west-central Montana (James C. Bergdahl pers. comm. 2008).
This species is known from two creeks near their confluences with the South Fork of the Payette River near Lowman in west-central Idaho (Bergdahl and Kavanaugh 2011: 81).
This species is known from a few localities in southeastern Washington and west-central Idaho (Bousquet 1985a: 259).
This species is known only from the type locality.
This species is known only from the type locality in northeastern Idaho.
This species is known only from several specimens collected at the type locality in northeastern Idaho.
Ten species in the temperate regions of North America, east of the Rocky Mountains.
Bousquet (1992b) published a preliminary key to the species.
This species is known only from the holotype.
This species is known only from two specimens collected in “Georgia.”
This species is known only from a few localities in west-central Georgia (Fattig 1949: 25), northern Florida (Bousquet 1992b: 511, 513) including the Panhandle (Wakulla County, CMNH), and “Louisiana” (Bousquet 1992b: 513).
This species is known only from several localities in Mississippi (Bousquet 1992b: 515; Drew A. Hildebrandt pers. comm. 2007) and western Alabama (Greene County, Foster F. Purrington pers. comm. 2010).
This species is known from a few localities from northern Georgia (Fattig 1949: 26, as
This species is known only from a few localities in Grayson County (CNC) in Texas.
This species ranges from southwestern Pennsylvania (Lindroth and Freitag 1969: 342) to “Iowa” (Jaques and Redlinger 1946: 297), south to east-central Louisiana (West Feliciana Parish, LSAM), northern Alabama (Löding 1945: 16; Madison County, CMNH), and northern Georgia (Fattig 1949: 25).
This species is known from a few localities in Oklahoma (Latimer and Marshall Counties, CMNH, CNC), eastern, central, and southern Texas (De Witt, Madison, Anderson, Lee, and Houston Counties, CMNH, CNC, MCZ; Riley 2011), and “Louisiana” (CNC).
This species is known from southwestern Florida (Collier County, Foster F. Purrington pers. comm. 2011) and from central and southern Mississippi (Will and Gill 2008: 121; Hancock and Leake Counties, CMNH, CNC) to southeastern Oklahoma (Pushmataha County, CNC), south at least to east-central Texas (Harris County, CNC).
This species is known from eastern Kansas (Lindroth and Freitag 1969: 341; Douglas County, CNC, UASM) to “Iowa” (Jaques and Redlinger 1946: 297) and southwestern Illinois (Saint Clair County, MCZ; Wickham 1895a: 185), south to northwestern Louisiana (Bossier Parish, LSAM; LeConte 1853a: 241) and south-central Texas (Comal County, CMNH). The record from “Florida” (Wickham 1895a: 185) needs confirmation (see Leng 1915: 576).
Two Palaearctic species (
Sasakawa and Kubota (2006) combined members of
This European species is adventive in North America on both coasts and is now found from Newfoundland (Lindroth 1955a: 87; Larson and Langor 1982: 592) to Vancouver Island (Lindroth 1966: 492), south to northwestern California (Del Norte County, James R. LaBonte pers. comm. 1992), northern Utah (Davis County, CMNH), northern Colorado (Miller and Peairs 2008: 34), South Dakota (Ellsbury et al. 1998: 621; French et al. 2004: 557; Larsen and Purrington 2010: 571), northeastern Iowa (Purrington and Larsen 1997: 50), east-central Ohio (Usis and MacLean 1998: 67), and central Pennsylvania (Capogreco 1989b: 4; Byers et al. 2001: 85). The first inventoried specimen collected on this continent was found in Nova Scotia in 1926 (Lindroth 1957c: 153) and in the west in Seattle in 1927 (Hatch 1933c: 120).
Two other subspecies of
Eleven species in the boreal and temperate regions of eastern North America (six species) and Japan (five species:
Nicolay and Weiss (1934) and Lindroth (1966) summarily reviewed the North American species. A thorough revision of this group is needed.
This subgenus is often combined with
This species occurs from New Brunswick (Bousquet 1987a: 124) to the Ontario Peninsula (Lindroth 1966: 495), south along the Appalachian Mountains to the Great Smokies in eastern Tennessee (Stockton 1954: Fig. 19; CNC) and to northern Georgia (Fattig 1949: 24). The record from southwestern Arkansas (Stockton 1954: 16) is probably in error.
This species is found from Newfoundland (Lindroth 1955a: 86) to eastern South Dakota (French et al. 2004: 557), north to the James Bay area in Quebec (Larochelle 1975: 86), south along the Appalachian Mountains to eastern Tennessee (Nicolay and Weiss 1934: 206, as
Until the status of this taxon is better defined, its distribution remains inadequately documented. It has been recorded from Indiana, Illinois, Iowa, Missouri (Casey 1918: 367), and South Carolina (Nicolay and Weiss 1934: 200, as
Lindroth (1966: 493) discussed the status of this taxon and at the time regarded it as an extreme, southern form of
This species is known from scattered localities from southwestern Quebec (Bousquet 1998: 105) to eastern North Dakota (Cass and Ransom Counties, CNC), north to southern Manitoba (Lindroth 1966: 495), south to southern South Dakota including the Black Hills (Kirk and Balsbaugh 1975: 23), northeastern Illinois (Purrington et al. 2002: 201), and northeastern New Jersey (FSCA). The record from Maryland (Erwin 1981b: 162), based on a single old specimen, needs confirmation.
The range of this species extends from northeastern New York (Notman 1928: 222) to southern Michigan (Eaton and Washtenaw Counties, UMAA), including southernmost Ontario (Lindroth 1966: 496), south to southwestern Indiana (Blatchley 1910: 93) and along the Appalachian Mountains to northern South Carolina (Ciegler 2000: 67) and central Alabama (Löding 1945: 16; Stockton 1954: Fig. 18). The records from eastern Kansas (Snow 1880: 78, as
This species ranges from Maine (Dearborn and Donahue 1993: 5) to central Iowa (O’Rourke et al. 2008: 126), south to “Louisiana” (Nicolay and Weiss 1934: 201) and southwestern Georgia (Fattig 1949: 24), west to northeastern Kansas (Popenoe 1877: 23). The record from “Minnesota” (Bousquet and Larochelle 1993: 174) needs confirmation. Fossil remnants from a Plio-Pleistocene sequence have been unearthed in northwestern Greenland (Böcher 1995: 26).
Eleven species in the arctic, subarctic, and boreal regions of North America (four species) and Asia (ten species, one of them extending to European Russia). Three species are Holarctic.
Lindroth (1966) reviewed the North American species. Budarin (1976) reviewed the world fauna, including two species (
This subgenus has been known for sometimes under the name
This Holarctic species is found in the arctic regions of eastern Siberia and northeastern China (Bousquet 2003d: 497) and on this continent from the Seward Peninsula in Alaska to northwestern Northwest Territories (Lindroth 1966: 528).
This Holarctic species ranges from the Yenisei River in Russia (Lindroth 1966: 530) to the Melville Peninsula in northern Nunavut, Canada [see Nielsen et al. 1987: Fig. 18c].
This species is found from Newfoundland (Lindroth 1955a: 87) to southeastern Yukon Territory (Lindroth 1966: 527), including northeastern British Columbia (CNC), north to northwestern Northwest Territories, south to central Alberta (Lindroth 1966: 527), northeastern Minnesota (Cook County, CNC), some mountains in New York (Notman 1928: 223), and “Massachusetts” (Wickham 1895a: 186). The record from Cuyahoga County in northern Ohio (Bubna 1902: 193) needs confirmation. Fossil remnants of this species, dated between about 11,700 and 18,100 years B.P., have been unearthed in northeastern Wisconsin, central Minnesota (Schwert 1992: 77), and central and southeastern Iowa (Schwert 1992: 77; Baker et al. 1986: 96).
This Holarctic species is found in the arctic regions from European Russia to the Melville Peninsula and Southampton Island in eastern Nunavut, Canada (Lindroth 1966: 528). Fossil remnants of this species, dated between about 16,700 and 18,100 years B.P., have been unearthed in southeastern Iowa (Baker et al. 1986: 96); others from a Plio-Pleistocene sequence have been found in northwestern Greenland (Böcher 1995: 28).
Eight species in North America (one Holarctic species) and Palaearctic Asia (eight species, one extending to European Russia).
According to Bousquet (1999: 145), this group is closely related and possibly paraphyletic in regard to
This Holarctic tundra species is found from the northern part of European Russia (Brinev and Shilenkov 2001) to northwestern Northwest Territories (Lindroth 1966: 531). Fossil remnants of this species, dated between about 16,700 and 18,100 years B.P., have been unearthed in southeastern Iowa (Baker et al. 1986: 96).
Five species in the temperate regions of eastern North America.
Lindroth (1966: 535) provided a key to all species, including also
Lindroth (1966: 534), followed recently by Lorenz (2005: 289), treated this group as a distinct genus. Structural characters of the adults and larvae clearly suggest that members of
This species ranges from east-central Iowa to North Carolina, including southern Michigan (Sadek 1982: 23), south to northern Florida (Peck and Thomas 1998: 20) and “Texas” (Schaupp 1882c: 41).
This species is known from scattered localities from Ohio (Purrington et al. 1989: 107; Usis and MacLean 1998: 67) and southwestern Pennsylvania (Horn 1880a: 139; Allegheny, Fayette, and Westmoreland Counties, CMNH) south to northern Georgia (Fattig 1949: 25).
In the
This species is found from eastern Maryland (Queen Annes County, Foster F. Purrington pers. comm. 2009) to southern Wisconsin (Sadek 1982: 34), including Pelee Island in southernmost Ontario (Lindroth 1966: 537), south to southern Louisiana (East Baton Rouge Parish, CNC) and the Florida Panhandle (Peck and Thomas 1998: 19).
This species occurs from central New York (Hajek et al. 2007: 880) to southeastern South Dakota (Kirk and Balsbaugh 1975: 23; Ellsbury et al. 1998: 621), north to southernmost Ontario (Bousquet 1987a: 125) and northeastern Michigan, south to northeastern Texas and northeastern Florida (Sadek 1982: 25, Fig. 16). The records from Quebec (Larochelle 1975: 31) and “New Mexico” (Bousquet and Larochelle 1993: 174) are based on mislabeled specimens or are in error.
The range of this species extends from “New York” (Wickham 1895a: 186) to Iowa (Wickham 1888: 82; Jaques and Redlinger 1946: 297), south to eastern Arkansas (Sadek 1982: 31), Alabama (Löding 1945: 16), Georgia (Fattig 1949: 25; House and All 1981: 195; CMNH, MCZ), and South Carolina (Ciegler 2000: 68).
One species in the Pacific Northwest.
The species was included in Lindroth’s (1966: 473) monograph.
This species is known from northern Washington to central Oregon (Foltz 2011).
One species in the boreal and temperate regions of North America.
The species was included in Lindroth’s (1966: 500) monograph.
This species has been included for a long time with the species of
This species is found from Newfoundland (Lindroth 1955a: 89-90) to the Okanagan Valley in south-central British Columbia, north to Fort Smith in southern Northwest Territories (Lindroth 1966: 500), south to northeastern Nevada (Elko County, CNC), central Colorado (Elias 1987: 632; Wickham 1902: 236), Missouri (Summers 1873: 134), and northern Georgia (Fattig 1949: 27).
Forty-two species in the arctic, subarctic, boreal, and temperate regions of North America (three species), Palaearctic Asia (34 species), and Europe (five species).
Lindroth (1966: 474-475, 406-497, as
This species is known from a few localities in Yukon Territory (Ball and Currie 1997: 451; CNC) and Alaska (Lindroth 1966: 475). The species is also cited, with a question mark, from the Magadan region in eastern Siberia (Budarin 1985: 15).
This species occurs from the neighborhood of New York City (Schaupp 1883b: 31; Ulster County, CMNH) to northeastern Ohio (Lee 1994: 60), south along the Appalachian Mountains to northern Georgia (Fattig 1949: 25; CNC) and northwestern South Carolina (Ciegler 2000: 67). The record from “Indiana” (Schrock 1985: 354) needs confirmation. The old records from Ontario and Quebec (see Lindroth 1966: 497) and Lake Superior (LeConte 1853a: 247) were based on misidentified or mislabeled specimens.
This species is known from scattered localities from Staten Island, New York (Smith 1910: 205) to northwestern Pennsylvania (Forest County, CMNH), south along the Appalachian Mountains to northeastern Georgia (Fattig 1949: 25). The record from “Michigan” (Wickham 1895a: 185) is in error.
Four species in the Appalachian region of eastern North America.
Darlington (1932) reviewed the species and provided a key for their identification.
This subspecies is known from the Black Mountains in western North Carolina (Darlington 1932: 162; Barr 1969: 72) and from northeastern Georgia near the border with South Carolina (Fattig 1949: 21). The record from “Tennessee” (Bousquet and Larochelle 1993: 171) needs confirmation.
This subspecies is endemic to the Great Smoky Mountains along the North Carolina-Tennessee border (Darlington 1932: 163).
This species is found along the Appalachian region from southern Quebec (Larochelle 1975: 101) to Kentucky (Powell County, CNC) and northeastern Georgia (Fattig 1949: 23). The records from northwestern Indiana (Wolcott and Montgomery 1933: 126, as
This species is known from the Roan, Grandfather, and Black Mountains in North Carolina (Barr 1969: 72) and northern Georgia (Fattig 1949: 23).
This species is apparently endemic to the Great Smoky and Unicoi Mountains along the North Carolina - Tennessee border (Barr 1969: 80).
Three species in the Appalachian region.
Barr (1971a) revised the species.
This subspecies occurs in the Appalachians from the Black Mountains in North Carolina to northern Georgia (Barr 1971a: 9). The record from “Kentucky” (Bousquet and Larochelle 1993: 176) refers to
This subspecies is known from a few localities in central Kentucky (Barr 1971a: 11) and southwestern Virginia (Hoffman 1998: 37).
This species is known from the Nantahalas to the Snowbird Mountains in Mason and Graham Counties, North Carolina (Barr 1971a: 12).
This species ranges from New Brunswick to southern Ontario (Lindroth 1966: 458), south along the Appalachian Mountains to northeastern Alabama (Löding 1945: 15) and northern Georgia (Fattig 1949: 21), including the higher mountains of Tennessee and North Carolina (Barr 1971a: 7). The records from “Iowa” (Bousquet and Larochelle 1993: 176) and southwestern Alabama (Löding 1945: 15) are likely in error.
Twenty-six species in western North America.
Hacker (1968) revised the species. Since then, four new species have been described by Will (2007), one species (
This species is restricted to the Coast Ranges in southwestern Oregon and northwestern California [see Hacker 1968: Fig. 35].
This subspecies is found from the Eel River in Humboldt County southwards to Mendocino County, California [see Hacker 1968: Fig. 35].
This subspecies is known only from Humboldt County, north of Eel River, in northwestern California [see Hacker 1968: Fig. 35] and from southwestern Oregon (Curry County, James R. LaBonte pers. comm. 2009).
This species is found in western California from near Mount Shasta in Siskiyou County south to the Monterey area in Monterey County [see Hacker 1968: Fig. 38].
This species is known only from the holotype collected in northwestern California.
This species is restricted to the Coast Ranges in Mendocino and Sonoma Counties, California [see Hacker 1968: Fig. 36].
This species ranges along the Coast Ranges from southern Oregon southwards to Humboldt County in northern California [see Hacker 1968: Fig. 37].
This species is found along the Pacific Coast of California south of the Eel River in Humboldt County to the mouth of the Navarro River in Mendocino County [see Hacker 1968: Fig. 37].
This species is, as far as known, restricted to a small area along the Pacific Coast from the Tomales Bay to the north side of the Pacific entrance of the San Francisco Bay [see Hacker 1968: Fig. 36].
This species is restricted to the Coast Ranges in southern Humboldt County and northern Mendocino County, California [see Hacker 1968: Fig. 37].
This species is found in southwestern Humboldt, southwestern Trinity, and northern Mendocino Counties in the Coast Ranges, northwestern California [see Hacker 1968: Fig. 37].
This species is known only from the holotype.
This species is restricted to a small area of the Sierra Nevada in Calaveras, El Dorado, and Tuolumne Counties, east-central California [see Hacker 1968: Fig. 39].
This species is restricted to the Sierra Nevada from the Yosemite National Park to near King Canyon National Park, California [see Hacker 1968: Fig. 39].
This species is restricted to a small area of the Sierra Nevada in Tuolumne County, California [see Hacker 1968: Fig. 39].
This taxon has been considered a subspecies of
This species is restricted to the Coast Ranges of California from Mendocino County to southern Sonoma County, east to western Napa County [see Hacker 1968: Fig. 38].
This species is known from two localities in Tehama and Butte Counties in northern California.
This species is found in northern Idaho and western Montana (Hacker 1968: 39).
This species is known from “Washington” to northern California along the Coast Ranges, from the Sierra Nevada, and from Reno in northwestern Nevada (Hacker 1968: 41). Hacker (1968: 41) implied that the species occurs in British Columbia but the record needs confirmation.
This species is known only from the type locality.
This species ranges from northwestern Oregon, as far east as the eastern slopes of the Coast Ranges, southwards to northwestern California [see Hacker 1968: Fig. 34].
This species is restricted to western Oregon, as far north as Mount Hood National Forest, and northwestern California [see Hacker 1968: Fig. 32]. The record from “Washington” in Bousquet and Larochelle (1993: 179) is based on two specimens from Yakima County which Hacker (1968: 11) regarded as mislabeled.
This subspecies ranges from southwestern British Columbia, including Vancouver Island (Lindroth 1966: 471), southwards to the Green Peter Mountain (Hacker 1968: 13) in the Oregon Cascades [see Hacker 1968: Fig. 33].
This subspecies is endemic to the Willamette Valley and surrounding foothills in western Oregon [see Hacker 1968: Fig. 33].
This species is known from four sites in Lincoln and Benton Counties in western Oregon (Brenner 2005).
This species is known from a few specimens collected in northern Idaho and along the western edge of Montana (Hacker 1968: 36; see Will and Gill 2008: Fig. 15).
This species is found in the Rocky Mountains in southern Alberta, southeastern British Columbia (Lindroth 1966: 470-471), northeastern Washington, northern Idaho (Hatch 1953: 112), and western Montana (Russell 1968: 59; Edwards 1975: 55; Hansen et al. 2009: 353). The record from “Alaska” (Erwin et al. 1977: 4.33; Bousquet and Larochelle 1993: 182) is probably in error.
This species is actually known only from the type locality (Hacker 1968: 37) in southern California.
Three species in the Pacific Northwest are currently recognized.
Hatch (1953) provided a key for the identification of the species.
This taxon is limited to the Coast Ranges in northwestern Oregon (James R. LaBonte pers. comm. 1994).
The name
This taxon is confined to the Coast Ranges in southwestern Oregon (James R. LaBonte pers. comm. 1994).
This species is confined to the Cascade Range in Washington and Oregon (James R. LaBonte pers. comm. 1994).
Sixty species in eastern (two species) and western (58 species) North America.
Casey (1913) reviewed most of the species then known (under
Bousquet (1999) combined members of
This species ranges from Cape Breton Island (Bousquet 1987d: 105) to northern Minnesota (Petrice et al. 2002: 9), south along the Appalachian Mountains at least to eastern Tennessee (Carter and Morgan Counties, CMNH) and northwestern South Carolina (Ciegler 2000: 66). The records from Georgia (Fattig 1949: 21), northern Alabama (Löding 1945: 15), and Iowa (Wickham 1911b: 6; King 1914: 321) need confirmation since they could refer to
This species has a similar distribution pattern than the preceding one ranging from Cape Breton Island (Bousquet 1987d: 105) to northern Minnesota (Petrice et al. 2002: 9), south to east-central Iowa (Iowa County, MCZ), Tennessee in the Great Smokies (CNC, MCZ), Georgia (Perrault 1973a: 37), and northwestern South Carolina (Ciegler 2000: 68). One old specimen labeled “La.” (MCZ) is probably mislabeled.
This species is found along the Pacific Coast from the Yakutat Bay in southeastern Alaska (Lindroth 1966: 465) to Mendocino County in California (Van Dyke 1919b: 5).
This species is known only from the lectotype collected in coastal California.
The lectotype of
I have seen this species from Marin and Fresno Counties in California (CNC); according to Van Dyke (1919b: 5), it extends south to Monterey County in California. The syntype studied by Lindroth (1966: 462) is not conspecific with the lectotype.
This species is known only from the lectotype collected in the Coast Ranges of California.
This species ranges from the southeastern coast of Alaska (Lindroth 1966: 466) to “California” (Dejean 1828: 327, as
This species is known from a few localities in southern Oregon (Hatch 1936: 702).
At present, this species is known only from the Coast Ranges in north and central California. The record from Utah (Knowlton and Wood 1947: 94) is no doubt in error.
This species is found along the Coast Ranges from Oregon (Lane County, CNC) to southern California (Fall 1901a: 44).
This species is restricted to the Coast Ranges and adjacent areas from the Gulf Coast of Alaska (Lindroth 1966: 460) to northwestern California (Notman 1929b: 222; Will and Gill 2008: 123).
As far as known, this species is found along the Coast Ranges in northern California.
This species is known from a few specimens collected in Klamath County, southern Oregon.
This species is found along the Rocky Mountains and their foothills in western Alberta, eastern British Columbia (Lindroth 1966: 466), northeastern Washington, northern Idaho (Hatch 1953: 109), and western Montana (Russell 1968: 58; Edwards 1975: 54; Will and Gill 2008: 123).
This species is known only from the holotype.
This species is known only from the San Bernardino Mountains in southern California.
This species is known only from two specimens collected in Mendocino County, California.
This species seems to be confined to the mountains of southern California (Fall 1901a: 44). The records from “Oregon,” “Nevada,” and “Nebraska” (Schaupp 1882c: 41) are probably in error.
This species is known from a few localities in the southern parts of the Sierra Nevada. The record from Baja California (Horn 1894: 308) needs confirmation.
This species is known only from the lectotype collected in northwestern California.
This species is known from the Coast Ranges and Sierra Nevada in central California.
This species is known only from the holotype.
This species is known only from the lectotype.
This species is known only from the type locality in western California.
This species is known only from Placer (Casey 1913: 124) and Tuolumne Counties (Dajoz 2007: 17) in the Sierra Nevada.
Casey (1913) based the description of this species on two specimens, one from Lake Tahoe, the other from Merced Creek. The specimen from Merced Creek was later (Casey 1918: 335) considered a distinct species which Casey named
This species is known only from the lectotype collected in west-central California.
Besides the lectotype, I have seen one specimen, probably of this species, from Mono County, California (CNC). The species was also reported from Zion National Park, Utah, by Tanner (1928: 270).
This species ranges from northwestern Montana (Russell 1968: 58) to Vancouver Island (Lindroth 1966: 464), south to the Coast Ranges in southwestern Oregon (Niwa and Peck 2002: 787) and Tuolumne County in the Sierra Nevada (Will and Gill 2008: 123). The record from southeastern Colorado (Wickham 1902: 235) is likely in error.
This species is known only from the Lake Tahoe area in the Sierra Nevada, California.
This species is known from the Cascade Range and Coast Ranges of western Oregon [see LaBonte 2006: Fig. 5].
This species occurs west of the Rocky Mountains from southern British Columbia, including Vancouver Island (Lindroth 1966: 463), to northern Oregon (Hatch 1953: 109, as
Prior to Lindroth (1966), this species was reported in the literature under the name
This species is known only from the lectotype collected in northern California.
This species is known from “Oregon” (LeConte 1853a: 239) and a few localities in the Sierra Nevada (Fall 1901a: 45).
This species occurs from the foothills of the Rocky Mountains in south-central Alberta to the Kootenay River drainage in southeastern British Columbia (Lindroth 1966: 463), south to the Sierra Nevada in California (Fall 1901a: 45; Papp 1978: 167), northern Arizona (Apache and Coconino Counties, CNC), and the Sangre de Cristo Mountains in New Mexico (Fall and Cockerell 1907: 158; Casey, 1913: 111, as
This species is known from northern Idaho and adjacent eastern Washington (Hatch 1953: 109).
This species is known only from Siskiyou County in northern California.
As far as known, this species seems to be restricted to southwestern California.
This name has been listed as a junior synonym of
This species is known so far only from southwestern California.
This very distinct species is still known only from the lectotype collected at Santa Barbara along the coast of California.
This species is known only from the type locality in southwestern California.
This species is known only from the original specimens collected in San Clemente Island, part of the Channel Islands, in the Pacific Ocean along the coast of southern California.
According to Fall (1901b: 44), this species is “common and widely diffuse” in southern California.
This species is known yet only from San Diego County in southwestern California.
This species is known only from the lectotype collected in southwestern California.
This large species is found from the Queen Charlotte Islands (Kavanaugh 1992: 70) south to southern California (Fall 1901a: 44, as
This species is known only from the lectotype collected in eastern California.
This species is found along the Coast Ranges in central California (Marin, San Mateo, and San Francisco Counties, Will and Gill 2008: 125, CNC). It was also reported from Santa Rosa Island in the Pacific Ocean (Fall 1901a: 44).
This species is found along the northern (Siskiyou County, CNC) and central parts of the Sierra Nevada (Will and Gill 2008: 118, 125).
In analyses using molecular data sequences, this species was consistently positioned as the sister-group to {
This species is known only along the coast in central Oregon [see Kavanaugh and LaBonte 2006: Fig. 14].
This species is found along the Pacific Coast from Vancouver Island to northern California (Lindroth 1966: 467).
This species is yet known only from the lectotype.
This name has been listed in synonymy with
This species is found along the Rocky Mountains from southern Montana (Hatch 1933a: 7) to northern New Mexico (Snow 1885: 67; Fall and Cockerell 1907: 157).
This very distinct species is known only from southwestern Oregon (Hatch 1951: 116; Niwa and Peck 2002: 787).
This species is known so far only from the Yosemite Valley in the Sierra Nevada, California.
This species is known for sure only from the Sierra Nevada, California. The record from “Oregon” (Schaupp 1882c: 40) needs confirmation.
This species ranges along the Pacific Coast from southern Alaska (Lindroth 1966: 462) to northern California (Fall 1901a: 45; Casey, 1913: 114, as
This species is known only from the type locality in southwestern California.
As far as known, the range of this species extends along the Coast Ranges of California from Humboldt County (Casey 1913: 114, as
This species is known only from the original specimens collected at the type locality in the Sierra Nevada.
This species seems to be restricted to the Coast Ranges in California. Fall (1901b: 44) statement that the species is “common in the Sierras and throughout the region [i.e., southern California] to the west” needs confirmation.
About 115 species in the arctic, subarctic, and boreal regions of North America (23 species) and the Palaearctic Region (about 100 species). Several species are Holarctic.
Ball (1966a) revised the North American species which he arrayed in three groups. Since then, two new species have been described, one (
Lorenz (2005: 275) listed
This Holarctic subspecies ranges from the Kola Peninsula in northern European Russia eastwards to Newfoundland (Ball 1966a: 119); the species is represented by isolated populations in the Gaspé Peninsula, Maine, New Hampshire, Vermont, New York, Michigan (Ball 1966a: 119-121), northeastern Minnesota (Cook County, CNC), and northern Wyoming (Big Horn County, FFPC) [see Ball and Currie 1997: Fig. 4]. The record from northern Colorado (Armin 1963: 217) is likely in error. Fossil remnants, older than 33,000 years B.P., have been unearthed in southwestern Ontario (Warner et al. 1988: 37); others from a Plio-Pleistocene sequence have been found in northwestern Greenland (Böcher 1995: 28).
The subspecies
This Holarctic species ranges from the Kuril and Commander Islands (Eremin 1998: 298) on the east coast of Asia to southern Yukon Territory, including Kodiak and Aleutian Islands, south to west-central British Columbia (Lemieux and Lindgren 2004: 562) [see Ball 1963: Fig. 3].
Lindroth (1966: 524) stated that the taxonomic status of this taxon as a distinct species from
This species is found from the western shore of Hudson Bay in Nunavut westwards to the coast of the Bering Sea in Alaska, southwards to northeastern British Columbia and northwestern Alberta (UASM) [see Nielsen et al. 1987: Fig. 18b].
The range of this Holarctic species extends from eastern Siberia (Ball 1966a: 131) eastwards to east-central Yukon Territory (Dempster Highway Mi 35, CNC), south to the Alaska Peninsula [see Ball 1963: Fig. 3].
This species ranges from the Labrador coast and Baffin Island to central Alaska, south at least to central British Columbia (Summit Lake, CNC); isolated on the Adirondack Mountains of New York, the White Mountains of New Hampshire, and the Shickshock Mountains in Gaspé Peninsula [see Ball 1963: Fig. 8; Nielsen et al. 1987: Fig. 17d; Ball and Currie 1997: Fig. 2]. Fossil remnants of this species, dated between about 16,700 and 18,100 years B.P., have been unearthed in southeastern Iowa (Baker et al. 1986: 96).
This species is known from two localities along the west coast of Alaska (Ball 1966a: 82).
This species ranges from northwestern Nunavut and central Northwest Territories westwards to northeastern Alaska (Ball 1962: 19). The record from McMurray in northern Alberta (Ball 1966a: 49), based upon a single specimen, has been rejected by Ball and Currie (1997: 476). Fossil remnants of this species, dated between about 16,700 and 20,530 years B.P., have been unearthed in southeastern and northeastern Iowa (Baker et al. 1986: 96; Schwert 1992: 77; Woodman et al. 1996: 17).
This subspecies is known only from the type locality in the arctic tundra and from the “Alpine interior Alaska” (Matthews 1974b: 1365).
This subspecies is known only from a few localities in the arctic from northwestern Northwest Territories (Ball 1966a: 66) to Prudhoe Bay in Alaska, including the Richardson Mountains in Yukon Territory (Ball and Currie 1997: 489).
This subspecies is known for sure only from Kotzebue on the Seward Peninsula (Matthews 1974b: 1365) and the type locality in northeastern Alaska. The record from “Yukon Territory” (Ball and Currie 1997: 452) could not be confirmed.
This subspecies is known from a few localities between Yukon River and Fairbanks in the Yukon-Tanana Highlands in Alaska (Ball and Currie 1997: 486).
This subspecies is known only from the type locality located at the mouth of the Anderson River along the northwestern coast of Northwest Territories (Ball and Currie 1997: 486).
This subspecies is known only from a few localities in central Alaska (Lindroth 1966: 515).
This species is found from the Hudson Bay coast in northern Ontario to northeastern British Columbia, north to central Northwest Territories [see Ball 1963: Fig. 9].
This species is known only from eastern and central Alaska [see Ball 1963: Fig. 8].
The range of this species extends from the Hudson Bay area in Nunavut to Kuskokwim Bay on the western coast of Alaska, south to northeastern British Columbia [see Ball 1963: Fig. 9] and northern Alberta (Birch Mountains, Gerald J. Hilchie pers. comm. 2009, determination of George E. Ball). The records from northern Colorado (Wickham 1902: 236; Armin 1963: 218) are likely in error.
This species is known from the Dempster Highway in Yukon Territory (UASM) to the Seward Peninsula in western Alaska [see Ball 1963: Fig. 8].
This Holarctic species is known from the Kamchatka Peninsula, several islands in the Bering Sea, mainland Alaska, and the Ogilvie Mountains in western Yukon Territory [see Ball 1963: Fig. 4].
This Holarctic species ranges from the Yenisei River in Siberia eastwards to the Mackenzie River delta in western Northwest Territories; isolated on the top of some mountains in New England and on the Shickshock Mountains of the Gaspé Peninsula, Quebec [see Ball 1963: Fig. 5]. The record from Wyoming (Lavigne 1977: 47) is in error. Fossil remnants of this species, dated between about 12,000 and 18,100 years B.P., have been unearthed in central and southeastern Iowa (Schwert 1992: 77; Baker et al. 1986: 96) and northeastern Pennsylvania (Barnosky et al. 1988: 178); others, older than 33,000 years B.P., have been found in southwestern Ontario (Warner et al. 1988: 37).
This species is known only from a few localities in the Seward Peninsula, western Alaska [see Ball 1963: Fig. 6].
This species is found from eastern Siberia to the Dempster Highway in Yukon Territory (UASM), including several islands of the Bering Sea [see Ball 1963: Fig. 4].
Matthews (1974b: 1365) remarked that some specimens of this species have anomalous male genitalia. He interpreted this condition as either
This species is known from the Norton Sound Inlet in western Alaska to the Coronation Gulf in northern Nunavut (Ball 1966a: 28-29).
This species is found along the Rocky Mountains from southern Alberta to Utah and Colorado (Ball 1966a: 26).
This Holarctic species is known in North America along the arctic coast from western Alaska to north-central Nunavut, and from Victoria and Banks Islands; in the Palaearctic Region the species is known only from the northern Bering Sea Coast (Ball 1966a: 47).
This species is known only from the Ogilvie Mountains in Yukon Territory.
This species ranges from northern Alaska to the western shores of the Hudson Bay in northern Manitoba and Southampton Island in Nunavut [see Ball 1963: Fig. 2; Nielsen et al. 1987: Fig. 18a]; isolated in eastern Ohio (Usis and MacLean 1998: 67). Fossil remnants of this species, dated between about 16,700 and 21,500 years B.P., have been unearthed in north-central Illinois (Garry et al. 1990: 394) and east-central Iowa (Baker et al. 1986: 96).
This species ranges from central Alaska to eastern Alberta, including western Northwest Territories, south to west-central Montana, northern Idaho, and northern Oregon [see Ball 1963: Fig. 2]. The records from northern Colorado (Packard 1877: 811; Wickham 1902: 236; Armin 1963: 218; Elias 1987: 632, as
This Holarctic species ranges from Novaya Zemlya in the Arctic Ocean off the coast of European Russia east to Franklin Bay on the coast of Northwest Territories, south in the Nearctic Region to central Northwest Territories and Umnak Island in the Aleutian Islands [see Ball 1963: Fig. 2]. Fossil remnants of this species, dated between about 16,700 and 20,530 years B.P., have been unearthed in northeastern and southeastern Iowa (Schwert 1992: 77; Baker et al. 1986: 96; Woodman et al. 1996: 17); others, older than 33,000 years B.P., have been found in southwestern Ontario (Warner et al. 1988: 37).
The subspecies
Forty-five species in the temperate and subtropical regions of North America with one species (
Freitag (1969) revised the species. Since the publication of his revision, a replacement name has been proposed for
This genus, certainly one of the most characteristic pterostichine elements in North America, could be closely related to
Eighteen species, most restricted to the Coastal and Gulf Plains.
This species is restricted to a small area from “Pennsylvania” (LeConte 1846b: 354) to North Carolina [see Freitag 1969: Fig. 126]. The record from Georgia (Fattig 1949: 24) needs confirmation.
The range of this species extends from West Sister Island (Ohio) in the Western Basin of Lake Erie (Will et al. 1995: 62) to northeastern Illinois, south to southern Alabama and northern Georgia [see Freitag 1969: Fig. 126]. The record from Missouri (Summers 1873: 134) needs confirmation.
This species is known from a small area in west-central and southern West Virginia (Roane and Raleigh Counties, CMNH), western Virginia, and North Carolina [see Freitag 1969: Fig. 126].
This species is found in southern Georgia and Florida including the Keys [see Freitag 1969: Fig. 129]. The record from northwestern South Carolina (Kirk 1970: 11) is probably in error.
This species is known only from a few specimens collected in “North Carolina,” northern South Carolina, and “Georgia” [see Freitag 1969: Fig. 129].
This species is known only from several specimens collected at the type locality in southern Alabama [see Freitag 1969: Fig. 129].
This species ranges from southern Georgia to southwestern Florida [see Freitag 1969: Fig. 125]. The record from Alabama (Löding 1945: 16) needs confirmation.
This species is known from a few localities in the western parts of the Florida Peninsula [see Freitag 1969: Fig. 125].
This species inhabits the Gulf Plain and southern Piedmont Plateau from South Carolina to the Florida Panhandle and “Louisiana” (LeConte 1853a: 231, as
This species is known only from Mobile County in southwestern Alabama (Freitag 1969: 118). The record from “Mississippi” (Bousquet and Larochelle 1993: 186) was based on a misidentified specimen (Drew A. Hildebrandt pers. comm. 2007).
This species is known only from southern Alabama, southern Mississippi (Drew A. Hildebrandt pers. comm. 2007), and east-central (West Feliciana Parish, LSAM) and southern Louisiana [see Freitag 1969: Fig. 128].
This species is known from southern Georgia and northern Florida [see Freitag 1969: Fig. 128].
This species is known from a few specimens collected in eastern Texas [see Freitag 1969: Fig. 128].
This species is found in a small area including eastern Tennessee, southwestern North Carolina, northwestern South Carolina, and northeastern Georgia [see Freitag 1969: Fig. 128].
This species is found along the Coastal Plain from South Carolina to central Mississippi [see Freitag 1969: Fig. 127].
This species ranges from southwestern Pennsylvania to southwestern Illinois (Union County, CNC), south to northern Alabama and northwestern Georgia [see Freitag 1969: Fig. 127].
This species is found in a small area from the District of Columbia south to southern South Carolina [see Freitag 1969: Fig. 127] including northeastern Georgia (LeConte 1853a: 230). The records from “New York” (Leng and Beutenmüller 1893: 138, as
This species is restricted to the Coastal Plain and southern Piedmont from South Carolina (Ciegler 2000: 71) to northeastern Alabama, south to the Florida Panhandle [see Freitag 1969: Fig. 127]. The record from North Carolina (Brimley 1938: 119) needs confirmation.
Twenty-seven species are known.
This species ranges from southern Georgia to southwestern Florida [see Freitag 1969: Fig. 131].
This species is endemic to a small area in the eastern part of central Florida [see Freitag 1969: Fig. 131].
This species is known from two localities in southeastern Texas [see Freitag 1969: Fig. 136].
This species is known from southeastern Oklahoma south to Comal County, Texas [see Freitag 1969: Fig. 136]. The record from “Arkansas” (Freitag 1969: 167) derives from the locality originally cited by Say (1823b: 145) which possibly refer to the Arkansas Territory. Prior to 1825, this territory included, besides Arkansas, all of the present state of Oklahoma. It could also refer to the Arkansas River which flows through Colorado, southern Kansas, northeastern Oklahoma, and Arkansas. The record from southeastern Kansas (Knaus 1907: 233) needs confirmation.
This species is known only from eastern Texas [see Freitag 1969: Fig. 136].
This species is known from the holotype and four specimens collected in Hinds County in southern Mississippi (Drew A. Hildebrandt pers. comm. 2009).
This species is known only from a few specimens collected in western Alabama and northeastern Mississippi [see Freitag 1969: Fig. 132].
This species is found from southwestern Pennsylvania to southeastern South Dakota, south to southern Colorado (Miller and Peairs 2008: 34; LeConte 1879d: 500; Wickham 1902: 236), Oklahoma, and central Arkansas [see Freitag 1969: Fig. 130]. The records from South Carolina (Kirk 1970: 11; Ciegler 2000: 70) are probably in error; that from “Kentucky” (Bousquet and Larochelle 1993: 188) needs confirmation.
This species is known from a few localities in eastern Oklahoma, southern Arkansas, and northwestern Louisiana (Natchitoches Parish, LSAM) [see Freitag 1969: Fig. 130].
This species is restricted to the Gulf Coastal Plain from southwestern Alabama to eastern Texas, north to southern Arkansas [see Freitag 1969: Fig. 132]. Freitag (1969: 142) suggested that the specimens labeled from Clay County in northeastern Kansas and Lawrence County in northeastern Arkansas are probably mislabeled.
This species is found only in eastern Texas (Freitag 1969: 143, Fig. 132).
This species is found along the Gulf Coastal Plain from eastern Texas and southern Arkansas to western Mississippi [see Freitag 1969: Fig. 132]. The record from northwestern South Carolina (Kirk 1970: 11) is in error.
This species is found west of the Appalachians Mountains from southwestern Pennsylvania to eastern Nebraska, north to southern Wisconsin and southeastern Minnesota, south to east-central Texas and southern Mississippi (Walthall County, Paul K. Lago pers. comm. 2009) [see Freitag 1969: Fig. 132].
This species ranges from southwestern Pennsylvania to the Mississippi River in western Illinois, south to southeastern Louisiana and southern Alabama [see Freitag 1969: Fig. 131]. The record from South Carolina (Kirk 1970: 11; Ciegler 2000: 70) is probably in error.
This species is found along the Appalachians, Piedmont Plateau, and Coastal Plain from Massachusetts to the Florida Panhandle and eastern Alabama [see Freitag 1969: Fig. 131]. The records from southeastern Louisiana (Summers 1874a: 80, as
This species is known from a few localities in southern Alabama and southern Mississippi [see Freitag 1969: Fig. 131].
This species ranges from northwestern Wisconsin to northeastern South Dakota, south to central Missouri [see Freitag 1969: Fig. 134]. The record from “North Dakota” (Bousquet and Larochelle 1993: 187) needs confirmation.
The range of this species extends from southern Pennsylvania and New Jersey to central Virginia [see Freitag 1969: Fig. 133]. The records from Ohio (Everly 1927: 155; Everly 1938: 141), “Michigan” (Leng 1920: 57), Indiana (Blatchley 1910: 101), North Carolina (Brimley 1938: 199), and Georgia (Fattig 1949: 22) are probably in error.
This species is known from a few localities in southeastern Minnesota, Iowa, and eastern South Dakota [see Freitag 1969: Fig. 134].
This species is restricted to Tennessee and northern Alabama [see Freitag 1969: Fig. 133].
This form has been treated as a subspecies of
This species is known only from Arkansas [see Freitag 1969: Fig. 133].
This subspecies is found west of the Mississippi River from southeastern Minnesota (Gandhi et al. 2005: 926), western Iowa and eastern Nebraska south to southern Kansas and central Missouri [see Freitag 1969: Fig. 133]. At least one specimen simply labeled from Arkansas is known (Freitag 1969: 150). The records from southern Wisconsin (Rauterberg 1885: 15) and northern Colorado (LeConte 1853a: 229, as
This subspecies ranges from southwestern Vermont (Bell and Nielsen 1978: 8) to northeastern Minnesota, including southernmost Ontario, south to northeastern Mississippi and eastern Tennessee [see Freitag 1969: Fig. 133]. The record from northwestern Missouri (Freitag 1969: 149) probably refers to
This species ranges from southeastern South Dakota and western Iowa south to central Texas, west to central Arizona [see Freitag 1969: Fig. 134].
This species ranges from southern Minnesota to southern Wyoming, south to the state of Durango and southeastern Texas [see Freitag 1969: Fig. 134]. The record from Missouri (Summers 1873: 134) needs confirmation.
This species is known from Texas, as far south as DeWitt County and west to El Paso County [see Freitag 1969: Fig. 135]. The record from “Oklahoma” (Bousquet and Larochelle 1993: 189) needs confirmation.
This taxon has been regarded as a subspecies of
This species is known from southern Oklahoma (Hatch and Ortenburger 1930: 11, as
This species inhabits the Great Plains from southwestern Minnesota (Gandhi et al. 2005: 926) to western South Dakota, south to southern New Mexico and southern Arkansas [see Freitag 1969: Fig. 135]. The record from “Arizona” (Bousquet and Larochelle 1993: 189) is probably in error; one specimen labeled from Hidalgo County in southeastern Texas is probably mislabeled (Freitag 1969: 163).
Eighteen species in Europe, with one species extending into Turkey, arrayed in two subgenera:
Sixteen species in Europe and Turkey, one of them adventive in the Nearctic Region.
The species in North America was covered in Lindroth’s (1969a: 1116) monograph.
This European species is adventive in North America where it is known from Sydney, Nova Scotia (Brown 1967: 87) and western Newfoundland (David W. Langor pers. comm. 1990). The first inventoried specimen collected on this continent was found in 1965 in Nova Scotia.
About 700 species arrayed in two subtribes:
About 595 species placed in one genus.
About 595 species (Hieke 2007) in the Nearctic (105 species, including ten adventive species), Neotropical (13 species in Middle America, only two of them endemic), Oriental (five species), Palaearctic (about 505 species, 16 of them Holarctic), and Afrotropical (about 12 species, nine of them endemic) Regions. These species are arrayed in 47 subgenera.
Almost all new state and province records listed here are based on specimens determined by Fritz Hieke.
Eighty-five species (Hieke 2007) in the arctic, subarctic, boreal, and temperate regions of North America (15 species of which three extend into Mexico), Asia (about 60 species), and Europe (13 species, all but possibly one shared with Asia). Six species are Holarctic and one is adventive in eastern North America.
There is no modern taxonomic revision of the species and such study would be useful. Lindroth (1968, as
This Holarctic species is found from Great Britain to the Pacific Coast of Siberia (Hieke 2003a: 558), and from the west coast of Alaska, including the Saint Lawrence Island, to northwestern Newfoundland, south to the James Bay area and to northern New Mexico along the Rocky Mountains [see Lindroth 1963b: Fig. 58]; isolated on the Shickshock Mountains in Gaspé Peninsula (Larochelle 1975: 42) and some mountains in New England (Lindroth 1966: 676). The record from “New York” (Notman 1928: 226) needs confirmation. According to Lindroth (1968: 676), this is the carabid occurring farthest north in North America. Fossil remnants of this species, dated between about 14,000 and 21,500 years B.P., have been unearthed in southeastern and central Iowa (Baker et al. 1986: 96; Schwert 1992: 78) and north-central Illinois (Garry et al. 1990: 394); others from interglacial deposits have been found in northwestern Greenland (Bennike 2000: 31).
This European species is adventive in North America where it is known from Newfoundland (Larson and Langor 1982: 593) and Saint Pierre and Miquelon (Lindroth 1955a: 95) to the Saint Lawrence Valley in Quebec (Larochelle 1975: 42), south to Boston Harbor, Massachusetts (Davidson et al. 2011: 512). The first inventoried specimen collected on this continent was found in Cape Breton Island, Nova Scotia in 1929 (Fall 1934: 171). The record from the Similkameen Valley in British Columbia (Smith et al. 2004: 96) was based on a misidentified specimen of
This western species occurs from southern Saskatchewan (MCZ) to south-central British Columbia (Lindroth 1968: 666), south to east-central California (Inyo County, CAS) and northwestern New Mexico (San Juan County, CNC).
This species ranges from eastern Siberia to the Hudson Bay coast in northern Manitoba (Lindroth 1968: 677).
This species ranges from the Gaspé Peninsula (Larochelle 1975: 43) to south-central British Columbia (Lindroth 1968: 669), south to the Sierra Nevada in California (Dajoz 2007: 16), central New Mexico (Bernalillo County, CMNH; Lindroth 1968: 668), the Texas Panhandle (Michels et al. 2010: 743), northwestern Arkansas (Carroll County, MCZ), and northeastern Ohio (Lake County, CMNH). The record from “West Virginia” (Bousquet and Larochelle 1993: 190) is in error; that from “Yukon Territory” (Ball and Currie 1997: 452) could not be confirmed.
This species is found in Kazakhstan, Mongolia, northern China, and across Siberia (Hieke 2003a: 559). In North America, it is known only from the Anderson River Delta area in northwestern Northwest Territories (Hieke 1994: 306).
The North American specimens differ from the Siberian ones in structural details and, according to Hieke (1994: 307), it is possible that they constitute a distinct subspecies.
This species ranges from Baja California Norte (CAS), southeastern California (Inyo and San Bernardino Counties, CAS, CMNH), and southern Nevada (Clark County, MCZ) east to northwestern Oklahoma (Texas County, CMNH), north to northern Colorado and “Utah” (Hayward 1908: 25, as
This Holarctic species ranges from northern Finland to the Pacific Coast, south to Mongolia and northeastern China (Hieke 2003a: 560) and from Alaska (Lindroth 1968: 679) to Newfoundland (Lindroth 1955a: 99), south to the upper peninsula of Michigan (Lindroth 1968: 678). The record from the White Mountains in New Hampshire (Wickham 1896b: 37) needs confirmation; those from “Vermont” (Hamilton 1894a: 10) and Colorado (Snow 1877: 17; Wickham 1902: 236; Armin 1963: 208, as
The range of this species extends from “Washington” (Hayward 1908: 23) to the Baja California Peninsula (Horn 1894: 309) and Arizona (Griffith 1900: 565; Hayward 1908: 23). The records from north-central Utah (Horn 1894: 309), “Colorado” (Wickham 1902: 236), and New Mexico (Fall and Cockerell 1907: 159) need confirmation.
This Holarctic species is known from northeastern Siberia and the Anderson River Delta in northwestern Northwest Territories (Hieke 1994: 311).
This species occurs from Nova Scotia (Larochelle and Larivière 1990a: 30, 34) to southwestern British Columbia, north to the coast of Nunavut and northern Alaska (Lindroth 1968: 671), south to northwestern Washington (Hatch 1953: 121), south-central Utah (Wayne County, CNC), southwestern Colorado (La Plata County, CNC), the Black Hills in southwestern South Dakota (Kirk and Balsbaugh 1975: 25), and “Michigan” (Garry A. Dunn pers. comm. 1986). The record from “Nebraska” (Bousquet and Larochelle 1993: 191) needs confirmation.
This species ranges from Nova Scotia (Lindroth 1968: 678) to “Wyoming” (Hayward 1908: 34), south to northeastern Colorado (Sedgwick County, Ken Karns pers. comm. 2009), northeastern Texas (Dallas County, MCZ), southwestern Alabama (Mobile County, USNM), and northwestern South Carolina (Ciegler 2000: 75). An old specimen simply labeled from “New Mexico” (MCZ) is known. The record from western Montana (Hatch 1933a: 8) needs confirmation.
This species ranges from southern Arizona (Cochise County, CNC) to western Texas (Dajoz 2007: 23; Brewster and Jeff Davis Counties, CNC, MCZ), south to the Federal District and central Veracruz in Mexico (Bates 1882a: 76).
The range of this species extends from southern Saskatchewan (CNC) to the Okanagan Valley in south-central British Columbia (Lindroth 1968: 668), south to northern Arizona (Coconino County, CMNH) and central New Mexico (Torrance County, CMNH; Milford et al. 2000: 21).
This circumpolar species is found from northern Finland to the Far East (Hieke 2003a: 561) and from the west coast of Alaska (Lindroth 1968: 673) to Newfoundland (Lindroth 1955a: 96-97), south to Nova Scotia (Lindroth 1968: 673), the Magdalen Islands, Gaspé Peninsula (Larochelle 1975: 48), southern Minnesota (Gandhi et al. 2005: 929), northern Colorado (LeConte, 1878a: 450, as
Thirty-five species (Hieke 2007) in the arctic, subarctic, boreal, and temperate regions of North America (12 species, of which two are adventive), Asia (26 species), and Europe (seven species, all shared with Asia). One species (
There is no modern taxonomic revision of the species. Lindroth (1968, as
This Palaearctic species is adventive in North America where it is known from Newfoundland (Lindroth 1955a: 100-101) and southern Labrador (Lindroth 1954d: 368) to the Kenai Peninsula in Alaska (Derek S. Sikes pers. comm. 2008), south to northern California (Trinity and Lassen Counties, CAS), west-central Nevada (Bechtel et al. 1983: 474), southern Colorado (Huerfano and Pueblo Counties, CMNH), Kansas (Trego County, CNC), and Virginia (Hoffman et al. 2006: 23; Clarke County, USNM). The first inventoried specimen collected on this continent was found in Quebec prior to 1865 (Couper 1865: 60, as
This species ranges from Newfoundland (Lindroth 1955a: 100) to southeastern British Columbia (Lindroth 1968: 690), south to southern Colorado (Wickham 1902: 236; Armin 1963: 200), southeastern Kansas (Knaus 1907: 233), Missouri (Summers 1873: 145), and southwestern North Carolina (Macon County, MCZ). The record from southeastern Louisiana (Summers 1874a: 80) is likely in error.
This species is known from a few localities in Yukon Territory and the Anderson River delta in northern Northwest Territories (Lindroth 1968: 687).
The range of this species extends from southern New Hampshire (Sullivan County, MCZ) to eastern South Dakota (Kirk and Balsbaugh 1975: 26; French et al. 2004: 557), south to northeastern Texas (Dallas County, MCZ), northeastern Georgia (Fattig 1949: 28; Clayton County, USNM), and northwestern South Carolina (Ciegler 2000: 74). The record from “Florida” (Bousquet and Larochelle 1993: 192) is probably in error.
This European species is adventive in North America where it is known only in eastern Canada from Newfoundland (Lindroth 1955a: 101) to the Gaspé Peninsula (Larochelle 1975: 44) and the north shore of the Saint Lawrence in Quebec (Brown 1932b: 200), including southern Labrador (Lindroth 1955a: 101). The first inventoried specimen collected on this continent was found in Newfoundland in 1905 (Lindroth 1955a: 101).
This species is found from the Yenisei River in western Siberia to the coast of Labrador [see Alfimov and Berman 2009: Fig. 1].
The range of this species extends from western Newfoundland (Lindroth 1955a: 102) to Vancouver Island (Lindroth 1968: 682), south to northwestern California (Humboldt County, USNM; Hayward 1908: 44), Arizona (Casey, 1918: 240, as
This species is known from a few localities in Labrador (CNC), northern Manitoba (CNC), northeastern Alberta (Andrew Lake, UASM), and Yukon Territory as far north as Swim Lakes (CNC), south to east-central Nevada (White Pine County, CMNH) and northern New Mexico (Hieke 1990: 234).
This species is known from Arizona (Coconino National Forest, CNC), northern New Mexico (Casey 1924: 48; Stehr 1949: 207, as
This species inhabits the Laurentian Highlands from the north shore of the Saint Lawrence Gulf (Larochelle 1975: 48) to the Great Slave Lake in the Northwest Territories (Lindroth 1968: 684), south to central Washington (Kittitas County, CMNH), southwestern Alberta (Indian Creek at 49º48’N - 114º07’W, CNC), northeastern Minnesota, and the upper peninsula of Michigan (Lindroth 1968: 684). The record from South Dakota (Kirk and Balsbaugh 1975: 26) was based on a small specimen of
This species is found along the Pacific Coast from central California (Lindroth 1968: 692) south to Guadalupe Island off the coast of Baja California (Casey 1918: 295, as
This species is known from “all the islands” (Fall 1901a: 45) off the coast of southern California.
One species in southern North America.
The species was redescribed in Hieke’s (1978: 290-292) revision of the subgenus
This species is known from McLennan (Horn 1892b: 27) and Dallas (Hieke 1978: 292) Counties in Texas and from southwestern Oklahoma (Kondratieff et al. 2005: 171).
Six species in the boreal and temperate regions of North America (one species), Asia (two species), Europe (three species, two shared with Asia), and northern Africa (two endemic species).
Hieke (1978) revised the species of this subgenus.
This species ranges from Newfoundland (Lindroth 1955a: 102) to southern Alaska (CNC), south to central California (Merced County, CAS), northern Arizona (Chen et al. 2006: 171; Apache and Coconino Counties, CMNH, CNC, UASM), northeastern Texas (Hieke 1978: 289), and northern Georgia (Fattig 1949: 28).
Thirty-two species (Hieke 2007) in the arctic (marginal), subarctic, boreal, and temperate regions of North America (11 species, two extending into northern Mexico), Asia (17 species), and Europe (ten species, many shared with Asia and one extending into northern Africa). One species (
Hieke (2001) revised the species. Two Asian species have been described subsequently.
This species ranges from northern Oregon to southern California and Sonora, east to western Texas and Chihuahua in Mexico (Hieke 2001: 128). The record from southwestern Utah (Tanner 1928: 270) needs confirmation.
This species is known from two localities, Manhattan in northeastern Kansas and Gatesville in central Texas (Hieke 2001: 136).
This species is found from western Maine (Oxford County, Ross T. Bell pers. comm. 2008) to southeastern Wyoming (Hieke 2001: 116), including southernmost Ontario (Lindroth 1968: 700), south to the Taos Mountains in northern New Mexico (LeConte 1876: 299; McKinley County, USNM), “Texas” (Horn 1892b: 32), southwestern Alabama (Hieke 2001: 116), northern Georgia (Fattig 1949: 29), and central South Carolina (Ciegler 2000: 74).
This western species ranges from southern Manitoba to southern British Columbia, south to southeastern California (San Bernardino County, David H. Kavanaugh pers. comm. 2008), southern Utah (Garfield County, CNC), south-central New Mexico, and northwestern Nebraska (Hieke 2001: 118).
This species ranges from Newfoundland to Alaska including Kodiak Island; it is also known from the Lake Superior region in northern Michigan (Hieke 2001: 113) and from northeastern New York (Essex County, CNC).
This form, listed as a synonym of
This species is known from a few localities in eastern Washington, northern Oregon, and west-central Idaho (Hieke 2001: 119).
This Holarctic species is found from western Northwest Territories to Alaska and from Kamtschatka to northern European Russia, south to Mongolia and northern China (Hieke 2001: 61).
This species ranges from northern California to north-central Colorado, south to southwestern New Mexico and southern Arizona (Hieke 2001: 126-127).
This form was considered a synonym of
This species ranges from the Kodiak Island and Kenai Peninsula in Alaska south to the Sierra Nevada in east-central California, northern Arizona, and northern New Mexico along the Rocky Mountains (Hieke 2001: 122-124).
This form was regarded as a synonym of
This subspecies ranges from northern California to northwestern Oklahoma (Cimarron County, CMNH), north to northern Colorado, south to Durango in Mexico (Hieke 2001: 135-136), including western Texas (Jeff Davis County, CMNH), and south-central California (Hieke 2001: 135).
Lindroth (1968: 697) listed
This subspecies ranges from northern Idaho and eastern Washington south at least to Monterey County in western California (Hieke 2001: 134).
This species is known from a few localities in Northwest Territories, Yukon Territory (CNC), Alaska, and from Manning Provincial Park in southern British Columbia (Hieke 2001: 110).
Twenty species (Hieke 2007) in northern North America (one Holarctic species) and eastern Asia.
Hieke (1999a) revised the species of this subgenus. Subsequently, ten new Chinese species were described.
This Holarctic species is found in Siberia east of the Lena River and in North America from Alaska to the Hudson Bay coast in Manitoba and Nunavut (Hieke 1999a: 347).
Forty species (Hieke 2007) in North America (12 species), Middle America (two species, one of them endemic,
There is no modern revision of the species of this subgenus and such work is much needed.
This European species is adventive in North America where it is known from Newfoundland (Lindroth 1955a: 105) to western Quebec (Larochelle 1975: 43), south to Massachusetts (Davidson et al. 2011: 513). The first inventoried specimen collected on this continent was found in Cape Breton Island in 1929 (Lindroth 1963a: Fig. 39).
This Holarctic species is found from northern Manitoba (CNC) to the Kenai Peninsula in Alaska (Lindroth 1968: 706), and in the Palaearctic Region from the Far East to Norway, south to Spain, Italy, Kazakhstan, and Mongolia (Hieke 2003a: 556). The records from British Columbia (Hamilton 1894a: 11; Jarrett and Scudder 2001: 379) need confirmation; those from “Washington,” “Colorado” (Hamilton 1894a: 11; Wickham 1902: 238), and Idaho (Horning and Barr 1970: 24, as
This subspecies ranges from Vancouver Island (Lindroth 1968: 694) to eastern South Dakota (Brookings County, USNM), south to Costa Rica (Hieke 1990: 221) and the Baja California Peninsula (Horn 1894: 309).
The subspecies
This species is known from southwestern Alberta (Waterton Lakes National Park, UASM), southern British Columbia (Jarrett and Scudder 2001: 379), Washington (Minsk and Hatch 1939: 215; Pierce County, UASM), eastern Oregon (Baker County, CNC), and northwestern Montana (Flathead County, UASM). There is one specimen, identified by Hieke, from Mayo in central Yukon Territory (CNC).
This form was considered a synonym of
This species is known from eastern Oregon (Baker County, USNM) to western South Dakota (Lawrence County, USNM), north to southeastern Montana (Powder River County, USNM), south to western Texas (Culberson County, CMNH) and central Arizona (Gila County, MCZ).
This species ranges from southern Alaska (Anchorage area, UASM) to southern Northwest Territories (CNC), south to northern Oregon (Hood River County, MCZ) and southern Colorado (Elias 1987: 633).
This widely distributed species ranges from Nova Scotia (Sable Island, CNC) to south-central British Columbia (Lindroth 1968: 708), south to southern California (San Diego County, CNC), southern New Mexico (Grant County, USNM; Wickham 1896c: 133; Fall and Cockerell 1907: 158), northern Oklahoma (French et al. 2001: 228; Alfalfa County, CMNH), central Mississippi (Casey 1918: 287, as
This northern species ranges from the coast of Labrador to central Alaska (Lindroth 1968: 707), south in the west to northern Idaho (Bonner County, CNC) and northwestern Montana (Edwards 1975: 56).
This species ranges from Nova Scotia (Majka et al. 2007: 9) to southwestern North Dakota (Tinerella 2003: 636), south to central Colorado (Summit County, UASM; Horn 1892b: 37; Wickham 1902: 238), northeastern Texas (Lamar County, Brian Raber pers. comm. 2010), northern Alabama (Madison County, CMNH, USNM), and northwestern South Carolina (Ciegler 2000: 75).
This species is found from Newfoundland (Lindroth 1955a: 104-105, as
This species is known from central and southern Texas (Horn 1892b: 38; Riley 2011; Bexar, Harris, and Lee Counties, MCZ) and central and southwestern New Mexico (Bernalillo and Luna Counties, CMNH). The record from southeastern Kansas (Knaus 1885: 58) needs confirmation.
This species is found along the Atlantic Coast from “New York” to northern Florida (Hieke 1993: 143).
Thirteen species in North America, of which two are also found in Mexico and three are Holarctic.
Lindroth (1968) covered all but four (
This species ranges from the Magdalen Islands in the Gulf of Saint Lawrence (CNC) to the Queen Charlotte Islands (Kavanaugh 1992: 75), south to southern California (San Bernardino County, CAS, MCZ), southern Arizona (Greenlee County, UASM), southern New Mexico (Otero County, MCZ), “Kansas” (Casey 1918: 252), central Michigan (Dunn 1982b: 37), and eastern New Brunswick (Webster and Bousquet 2008: 18).
This circumpolar species is found over most of the boreal-alpine region in the Palaearctic Region and in North America from Alaska (Lindroth 1968: 717) to Newfoundland (Lindroth 1955a: 107), south to New England (Lindroth 1968: 716), northern Wisconsin (Messer 2010: 38), southern Colorado (Mineral County, UASM; Lindroth 1968: 716), northern Utah (Salt Lake County, CMNH), and Mount Rainier in western Washington (Lindroth 1968: 717). Several state records (e.g., CA, IA, MI, MN, ND, NM, OR) listed in Bousquet and Larochelle (1993: 196) and taken from earlier authors probably refer to other species or need confirmation.
This species ranges from eastern Manitoba to southwestern British Columbia, north at least to the Magunday River in southern Yukon Territory (Lindroth 1968: 714), south to southeastern California (San Bernardino County, CAS), southern Arizona (Maricopa County, UASM), southern New Mexico (Grant County, USNM; LeConte 1855: 353), and Durango in Mexico (UASM). The record from Wisconsin (Rauterberg 1885: 17) is probably in error; that from “Ontario” (Bousquet and Larochelle 1993: 196), based on a specimen from Nipigon Bay (CMNH) determined by Hieke, needs confirmation as the specimen could be mislabeled or could be a stray.
This Holarctic species ranges from Finland to the Kamtschatka Peninsula (Hieke 2003a: 556) and from Alaska to northwestern Northwest Territories (Lindroth 1968: 716). The records from British Columbia (Jarrett and Scudder 2001: 380) and west-central Montana (Hansen et al. 2009: 353) need confirmation.
This boreal species is found from Newfoundland to southwestern British Columbia, north to east-central Alaska (Lindroth 1968: 717), south to southwestern California (Ventura County, MCZ), northeastern New Mexico (Snow 1885: 67; Las Vegas, CMNH) along the Rocky Mountains, the Black Hills in western South Dakota (Lawrence County, USNM), northeastern Minnesota along Lake Superior (Casey 1918: 312, as
This species ranges from northern Arizona (Coconino County, USNM) and New Mexico (San Miguel County, USNM) south at least to the Isthmus of Tehuantepec (Bates 1882a: 77).
This species is known only from the Sierra Nevada in California (Casey 1918: 253; Alpine, Amador, Eldorado, Placer, Plumas, and Shasta Counties, CAS, MCZ, USNM).
This species ranges from Newfoundland (Lindroth 1955a: 107) to the Seward Peninsula in western Alaska (Lindroth 1968: 715), south to “California” (Lindroth 1968: 715), central Colorado (Elias 1987: 633; Casey 1918: 252, as
This species is known from Andrew Lake in northeastern Alberta to south-central British Columbia, south to southeastern California, southeastern Arizona, and northeastern New Mexico (Hieke 2002: 660).
This species is known only from a few specimens collected in Colfax and Catron Counties, New Mexico (Hieke 1997: 250).
This species is known from the Rocky Mountains in northern Arizona (Hieke 1990: 223) and from the states of Durango (Bates 1891a: 248, as
This species is known from the Hudson Bay region in northeastern Manitoba, from northern British Columbia (CNC), and from the Northwest Territories to the Seward Peninsula in Alaska. In the Palaearctic Region, the species ranges from eastern Siberia to the Taimyr Peninsula, south at least to the Yakutsk area (Hieke 2002: 668).
Eighty-seven species (Hieke 2007) in North America (29 species, of which six are adventive), Middle America (four species of which one,
No taxonomic revision has been published for the species of this group and such study is much needed. Hieke (2000) revised the six species of the
The range of this western species extends from western Montana (Russell 1968: 61; Hansen et al. 2009: 353) to Vancouver Island, north to the Prince Rupert area in British Columbia (Lindroth 1968: 711), south to Baja California Norte (CAS). The records from northern Colorado (Armin 1963: 201) and South Dakota (Kirk and Balsbaugh 1975: 26) need confirmation.
This species ranges from “Maine” to “Montana” (Hieke 2003b: 207), south to southern Colorado (Elias 1987: 633), southern Texas (Bastrop and Kerr Counties, CNC), and the Florida Panhandle (Okaloosa County, CNC); also recorded from “Oregon” (Hieke 2003b: 207).
This species is found from Prince Edward Island (King County, CNC) and Nova Scotia (NSNH) to southwestern British Columbia (Creston area, UASM), south to northern Idaho (Hatten et al. 2007: 359), northern Colorado (Miller and Peairs 2008: 34; Larimer County, UASM), southern Oklahoma (Elliott et al. 2006: 126), southwestern Alabama (Mobile County, USNM), and North Carolina (Hieke 2003b: 205). Several state records (e.g., AR, DC, DE, GA, MS, NE, SC, UT) reported in Bousquet and Larochelle (1993: 195) refer to other species of the group or need confirmation.
This species is yet known only from two specimens, both holotypes.
This form was considered a synonym of
This species ranges from Nova Scotia (Hieke 2003b: 206) to the Rocky Mountains in Alberta (CNC), south to “Colorado,” “Illinois,” and “New Jersey” (Hieke 2003b: 206).
This form was considered a synonym of
This species is found from southern Manitoba (Casey 1924: 63, as
This form was considered a synonym of
This species is found east of the Rocky Mountains from Maine to at least southern Wisconsin, including southern Quebec and the Ontario Peninsula, south to southern Texas (Hieke 2000: 69), southeastern Louisiana (East Baton Rouge Parish, Igor M. Sokolov pers. comm. 2009), and southern Georgia (Torres and Ruberson 2006: 31). One specimen is known also from Nuevo León in Mexico and four from Costa Rica (Hieke 2000: 69). Several old records listed by Bousquet and Larochelle (1993: 196) (e.g., CO, CT, DE, FL, MB, MN, MS, NB, NF, NM, NS, PE, RI, SD, SK, VT) refer to
This Holarctic species ranges from Newfoundland to the Gulf Coast of Alaska, including Kodiak Island (Lindroth 1968: 731), south to southern California (Dajoz 2007: 20; San Diego County, CNC), northern New Mexico (Casey 1918: 297, as
This species ranges from Newfoundland to southern Manitoba, south to northeastern Colorado, central Missouri, and North Carolina (Hieke 2000: 72-74). One specimen simply labeled from Texas is known (Hieke 2000: 69).
Lindroth (1968: 728) regarded this form as a synonym of
This European species is adventive in North America where it is known in the east from Nova Scotia (Majka et al. 2006: 605) to southern Wisconsin (Messer 2010: 38), south to Virginia (Hieke 2000: 82). The first inventoried specimen collected on the east coast was found in Massachusetts in 1925. The species is also adventive in the western part of North America where it is known from southeastern British Columbia and Alberta (Hieke 2000: 82). The first inventoried specimen collected on the west coast was found in southeastern British Columbia in 1936 (see Hieke 2000: 82). One specimen simply labeled from Iowa is known (Hieke 2000: 82). The record from “Cleveland, Or.” (Hieke 2000: 82) obviously refers to Ohio.
The range of this species extends from southern Arizona and western Texas south to Guatemala (Hieke 1993: 110).
This species ranges from southern Quebec to southeastern Alberta, south to southern Arizona, southern Texas, west-central Alabama, and North Carolina (Hieke 2000: 76-79).
Lindroth (1968: 730) regarded this form as a synonym of
This European species is adventive in North America where it is known in the east from Newfoundland (Lindroth 1955a: 111) to southwestern Manitoba (Stjernberg 2011: 71), south to northeastern Oklahoma (Hieke 1990: 203), northern Louisiana (Morehouse and West Carroll Parishes, Igor M. Sokolov pers. comm. 2009), and the Florida Panhandle (Okaloosa County, CNC) and in the west from southern British Columbia (Jarrett and Scudder 2001: 378; CNC) to southeastern Alberta (CNC), south to northern Colorado, north-central Arizona, and the San Francisco Bay area in west-central California [see Hieke 1990: Fig. 23]. The first inventoried specimen collected in the eastern part of this continent was found in Brooklyn, New York, in 1904 and in the western part in San Francisco, California, in 1941 (see Hieke 1990: 202, 204). This species was recorded from “Amérique septentrionale” by Dejean (1828: 466), under the name
This species ranges from Newfoundland (Lindroth 1955a: 108) to west-central Northwest Territories, south to northern Alberta (Lindroth 1968: 723), east-central South Dakota (Kirk and Balsbaugh 1975: 27), central Iowa (O’Rourke et al. 2008: 126; Larsen et al. 2003: 292), and northeastern New York (Essex and Hamilton Counties, USNM); apparently isolated in central Colorado (Elias 1987: 633). The record from “Yukon Territory” (Ball and Currie 1997: 453) could not be confirmed.
This European species is adventive in North America where it is known from southern British Columbia (Jarrett and Scudder 2001: 379) to southern California (Los Angeles County, CMNH). The first inventoried specimen collected in that area was found in southwestern Washington in 1929 (Hatch and Kincaid 1958: 5). The species has been recorded also from the Washington D.C. area since 1964 (Hieke 1990: 205) and has been recently collected in Westchester County in southeastern New York (Peter W. Messer pers. comm. 2011).
This species ranges from “New Hampshire” (Hayward 1908: 54) to southeastern South Dakota (Kirk and Balsbaugh 1975: 27), south to Oklahoma (Alfalfa and Logan Counties, CMNH, USNM), northeastern Georgia (Clarke County, USNM), and central South Carolina (Ciegler 2000: 74). One old specimen labeled “Tex” (USNM) is known.
This species ranges from southern Manitoba to south-central British Columbia (Lindroth 1968: 724-725), south to east-central Washington (Casey 1924: 60, as
This European species is adventive in North America where it is known from the Maritime Provinces (Bousquet 1992a: 504; Majka 2005: 534) and Connecticut (Majka 2005: 534) [see Majka 2005: Fig. 1]. The record from the Gaspé Peninsula, Quebec (Majka 2005: 534), is based on a misidentified specimen of
This species ranges from northwestern Montana (Russell 1968: 60) to Vancouver Island (Lindroth 1968: 728), south to southern California (San Diego County, CAS, CNC) and northern Utah (Davis, Salt Lake, and Utah Counties, CMNH).
This species is found from southern Manitoba to south-central British Columbia, north to central Alaska (Lindroth 1968: 726), south to the Sierra Nevada in east-central California (Tuolumne County, MCZ), central Arizona (Gila County, MCZ), central New Mexico (Fall and Cockerell 1907: 158), southern Kansas (Knaus 1905a: 218; Casey 1918: 311), and northern Indiana (Blatchley 1910: 107, as
The range of this species extends from Nova Scotia (Majka et al. 2007: 9) to central British Columbia (Lindroth 1968: 735), south to southeastern Arizona (Greenlee County, MCZ), southern New Mexico (Grant County, USNM), western Texas (Jeff Davis County, Ken Karns pers. comm. 2009), and southern New Jersey (Cape May County, MCZ). The record from North Carolina (Brimley 1938: 122, as
This species is found along and west of the Rocky Mountains from southwestern Alberta to south-central British Columbia, south to “Utah” and northern Colorado (Hieke 2002: 642).
This European species is adventive in North America where it is known only from a few localities in eastern Newfoundland (Bousquet 1987a: 128-129). The first inventoried specimen collected on this continent was found in 1971.
This species is known only from two specimens collected in Maryland [not Pennsylvania as stated by Hieke (2002: 643)] and North Carolina (Hieke 2002: 643).
This European species is adventive in North America where it is known in the east from Newfoundland (Lindroth 1955a: 111) to southern Saskatchewan (Ronald R. Hooper pers. comm. 2007), south to central Oklahoma (Grady County, Robert L. Davidson pers. comm. 2012), northwestern Mississippi (Panola County, CMNH) and the Florida Panhandle (Okaloosa County, CNC) [see Hieke 1990: Fig. 25]. The first inventoried specimen collected on the east coast was found in Rhode Island in 1901 (Hieke 1990: 207). The species is also adventive in the western parts of North America where it ranges from southeastern Alaska to central California, east to western Montana [see Hieke 1990: Fig. 25]. The first inventoried specimen collected on the west coast was found in western Washington in 1913 (Hatch 1953: 26).
This circumpolar species ranges from eastern Siberia to Ireland, south to Spain, Italy, Mongolia, and northern China (Hieke 2003a: 550) and from Alaska (Lindroth 1968: 721) to Newfoundland (Lindroth 1955a: 109), south to northeastern West Virginia (Randolph County, CMNH), east-central Ohio (Usis and MacLean 1998: 67), southern New Mexico (Otero County, CNC), and southwestern Oregon (Niwa and Peck 2002: 787).
This species is known from a few specimens collected in Baker and Klamath Counties in Oregon and Marin and El Dorado Counties in California (Hieke 2002: 654).
This species ranges from northern Washington south to the Mexican border in southern California, east to eastern Nevada (Hieke 1993: 114). One specimen simply labeled from Mexico is known (Hieke 1993: 114).
This species is, as far as known, restricted to southern British Columbia, including Vancouver Island (Lindroth 1968: 722), northern Washington, northern Idaho (Hatch 1953: 124), and northwestern Montana (Russell 1968: 60).
Twenty-one species (Hieke 2007) in the Palaearctic Region (including the Himalayas and northern Africa), one of them (
Hieke (2006) revised the species of this subgenus.
This Holarctic species is widely distributed in the Nearctic Region from Newfoundland (Lindroth 1955a: 103) to Alaska, including the Kodiak, Aleutian and Pribilof Islands (Lindroth 1968: 696), south to southern California (Fall 1901a: 45, as
The subspecies
Twenty-five species (Hieke 2007) in North America (nine species, all endemic), Asia (six species, only one endemic), Europe (13 species, six endemic), and northern Africa (four species, two endemic).
Lindroth (1968: 735-741, as
This species ranges from southern Quebec to southeastern British Columbia, south to southeastern Wyoming, southeastern Texas (Fort Bend and Harris Counties, Brian Raber pers. comm. 2010), east-central Louisiana (West Feliciana Parish, Igor M. Sokolov pers. comm. 2009), southern Mississippi (Hinds County, Drew A. Hildebrandt pers. comm. 2010), northern Alabama (Lawrence County, UASM), and South Carolina (Hieke 2000: 49-51). The records from “St. Louis, Montana” and “Cleveland, Oregon” listed by Hieke (2000: 51) refers to Missouri and Ohio respectively. The records from “New Brunswick” and “Prince Edward Island” (Bousquet and Larochelle (1993: 199) refer to other species; that from west-central Colorado (Wickham 1902: 237) needs confirmation.
This species ranges from New Brunswick to South Dakota, south to southern Pennsylvania (Hieke 2000: 55-57) and northeastern West Virginia (Randolph County, CMNH).
This species is known from a few localities in central and southern Texas (Hieke 2000: 58).
This species ranges from Cape Breton Island to southern Manitoba, south to northern Arizona, northeastern Colorado, northern Illinois, and northeastern Virginia (Hieke 2000: 53-55). The record “Cleveland, Oregon” (Hieke 2000: 55) refers to Ohio.
This form has been treated as a synonym of
This species occurs along the Pacific Coast from central Oregon (Lincoln County, CNC) to Monterey County in California (Hieke 1990: 196).
This species is known from southern Oregon (Lake County, CNC) and several localities in northern and central California (Hieke 1990: 198).
This species is found from southwestern British Columbia, including Vancouver Island (Lindroth 1968: 740), to southern California (Hayward 1908: 46; Moore 1937: 10; CAS) and “Arizona” (Casey 1918: 318, as
The range of this species extends from Cape Breton Island (Lindroth 1954c: 305) to Vancouver Island (Lindroth 1968: 739), north to Fort Smith in southern Northwest Territories (Bousquet 1987a: 130), south to western Washington (Hatch 1953: 129), southwestern Colorado (Montezuma County, CNC), Iowa (Hayward 1908: 46; Wickham 1911b: 6; Dickinson County, USNM), and Virginia (Casey 1918: 314). The records from “Nevada,” “Tennessee” (Bousquet and Larochelle 1993: 200), and Georgia (J.E. LeConte 1849: 26; Fattig 1949: 30) need confirmation.
This species ranges from southern Manitoba to Vancouver Island (Lindroth 1968: 740), south to southern California (Fall 1901a: 45; Moore 1937: 10) and southwestern Colorado (Wickham 1902: 237), east to northwestern South Dakota (Kirk and Balsbaugh 1975: 27).
Worldwide, with about 300 species in 32 genera (Lorenz 2005: 324-327). The Northern Hemisphere is represented by about 45 species (only 14.5% of the world fauna) and the Western Hemisphere by 55 species (about 18.5%). Seventeen species occur in North America.
Bousquet (1996a) revised and provided a key to the Nearctic, Mexican, and West Indian species.
One species in temperate North America.
This species is known from a few scattered localities in North Carolina (Brimley 1938: 127), Rabun County in northeastern Georgia (Fattig 1949: 45), southwestern Alabama (Washington County, Paul K. Lago pers. comm. 2009), northeastern Mississippi (Tishomingo County, Drew A. Hildebrandt pers. comm. 2008), southern Louisiana (Bousquet 1996a: 453), central Arkansas (Pulaski County, CMNH), eastern Missouri (Anonymous 2007), east-central Illinois (Coles County, Robert L. Davidson pers. comm. 2012), eastern Oklahoma (Bousquet 1996a: 453), and eastern Texas (Sabine County, Brian Raber pers. comm. 2010; Riley 2011).
One species in temperate North America.
This species is restricted to the Coastal Plain ranging from the District of Columbia to the Florida Panhandle, west to southeastern Mississippi (Drew A. Hildebrandt pers. comm. 2007) [see Bousquet 1996a: map 1].
Six species (Bousquet 1996a: 456), possibly eight (Lorenz 2005: 324), in the temperate and tropical areas of the Nearctic (two species), Neotropical, Australian, and Oriental Regions.
This species ranges from Massachusetts to central Kansas, south to southeastern Texas and southern Florida [see Bousquet 1996a: map 2]; also recorded from the Bahamas (Turnbow and Thomas 2008: 10).
This species ranges from eastern Texas south to Honduras; it also occurs in the Bahamas and Cuba [see Bousquet 1996a: 459, map 3].
Ten species in the temperate and tropical areas of the Nearctic (two species), Neotropical, and Australian (one species in the Malay Archipelago) Regions.
This species ranges from southeastern Texas (Bousquet 1996a: 465; map 5) south to Columbia (Martínez 2003: 17).
This species is known from the Florida Peninsula and several islands of the West Indies [see Bousquet 1996a: map 4].
Three species in temperate regions of North America (one species) and eastern Asia (two species).
This species has a disjunct distribution over eastern North America. In the north, it ranges from Nova Scotia (Majka et al. 2007: 11) to southeastern Manitoba, south to Nebraska, northwestern Tennessee (Obion County, CMNH), and Maryland; in the south, it occurs along the Coastal Plain from Georgia (Fattig 1949: 45) to southern Florida, west to southern Louisiana [see Bousquet 1996a: map 6]. The species could reach the Rio Grande since Summers (1874b: 135) reported seeing one specimen “from Mexico, near the Rio Grande.”
About 50 species in temperate and tropical areas of the Nearctic (four species), Palaearctic (11 species), Oriental, Afrotropical (one species), and Australian Regions.
This species ranges from southern Maine to east-central Minnesota (Kamal J.K. Gandhi pers. comm. 2008), south to southeastern Texas and the Florida Peninsula; also known from one locality in Cuba [see Bousquet 1996a: map 7]. The record from southeastern Wyoming (Lavigne 1977: 47) is probably based on a mislabeled specimen or a stray.
This species is restricted to the Coastal Plain ranging from Maryland to central Florida, west to central Louisiana [see Bousquet 1996a: map 9].
This species ranges from southern New Hampshire to northwestern Arkansas, north to southernmost Ontario and the central part of the lower peninsula of Michigan, south to eastern Texas and the Florida Panhandle [see Bousquet 1996a: map 10]. The record from “Iowa” (Bousquet and Larochelle 1993: 200) needs confirmation.
This species occurs from southern Maine and southern Quebec to western Minnesota, south to east-central Texas (Riley 2011), southwestern Louisiana and southern Florida [see Bousquet 1996a: map 8].
About 30 species in temperate and tropical areas of the Nearctic (six species) and Neotropical Regions, including the West Indies. The species are currently arrayed in two subgenera:
Ten species, ranging from southern Texas to central Argentina and including the West Indies, belong to this subgenus.
This species ranges from southern Texas to southern Mexico [see Bousquet 1996a: map 11]; it is also known from the Bahamas, Cuba, Cayman Islands, Trinidad and Tobago, and French Guiana (Bousquet 1996a: 482-483). The record from southeastern Louisiana (Summers 1874a: 80, as
Twenty species in temperate and tropical areas of the Nearctic (five species) and Neotropical (19 species) Regions, including the West Indies.
This species ranges from southeastern Wisconsin (Messer 2010: 39) and southernmost Ontario south to southern Florida and the Rio Grande in southern Texas, west along the Rio Grande to central New Mexico [see Bousquet 1996a: map 15]. The species is not yet know from Mexico but it certainly occurs at least along the states bordering Texas. The record from “Pennsylvania” (Bousquet and Larochelle 1993: 201) needs confirmation.
This species ranges from Long Island, New York, to south-central Oklahoma, south to Nicaragua (Bates 1882a: 46) and southern Florida, west to the Mexican state of Nayarit; also known from several islands in the Bahamas and the Greater Antilles [see Bousquet 1996a: map 12].
This species ranges from southwestern California to eastern Arizona, south along the Gulf of California to Nayarit [see Bousquet 1996a: map 17]. A few specimens simply labeled from New Mexico and Texas are known.
This species ranges from New Jersey to southern Wisconsin, south through southeastern Kansas (Knaus 1903: 188) and eastern and central Texas to southern Mexico, and to the Florida Keys, west to the Pacific Coast in the Mexican state of Colima; also recorded from the Bahamas (Turnbow and Thomas 2008: 15) and Cuba [see Bousquet 1996a: map 13]. The record from “New Mexico” (Bousquet and Larochelle 1993: 201, as
This species ranges from southern Texas south to Argentina (Brullé 1837: 32, as
Worldwide, with about 270 species in 19 genera (Lorenz 2005: 319-323, as
Fourteen species in temperate and tropical areas of the Nearctic (three species), Neotropical (four species), and Palaearctic (eight species) Regions arrayed in two subgenera:
Ogueta (1966a) reviewed the Western Hemisphere species and provided a key for their identification.
Western Hemisphere, with six species in the Nearctic (three species) and Neotropical (four species) Regions.
This species ranges from Long Island, New York (Notman 1928: 213; MCZ), to southern Florida (Peck and Thomas 1998: 20), west to eastern Texas along the Gulf Coast (Snow 1906a: 141; Galveston County, USNM); also recorded from northern Indiana (Blatchley 1910: 65; Schrock 1985: 352) and along the Ohio River in southwestern Ohio (Dury 1902: 111). The record from “Pennsylvania” (Bousquet and Larochelle 1993: 202) needs confirmation.
This species is found from Massachusetts (Harris in Scudder 1869: 264) and Connecticut (Krinsky and Oliver 2001: 136) to western South Dakota (Larsen and Purrington 2010: 571), including southern Ontario (Lindroth 1969a: 969), south to southeastern Texas (San Patricio County, CNC) and southern Florida (Peck and Thomas 1998: 20); also recorded from northeastern New Mexico (Fall and Cockerell 1907: 157).
This species ranges from central and southern Arizona (Cochise, Santa Cruz, and Yavapai Counties, CMNH, CNC; LeConte 1879c: 60) and western and south-central Texas (Presidio and Bexar Counties, CMNH, MCZ; LeConte 1879c: 60), south at least to central Veracruz and the Federal District in Mexico (Ogueta 1966a: 12); also found in the Baja California Peninsula (Horn 1894: 307).
Two species in temperate and tropical areas of the Nearctic (one species) and Neotropical (two species in Mexico) Regions.
Van Dyke (1927b: 93) discussed the structural differences between the two species.
This species ranges from southwestern Tennessee (Hardeman County, CMNH) to southwestern Arizona (Maricopa County, USNM), north to southeastern Colorado (Michels et al. 2008; Las Animas County, Robert L. Davidson pers. comm. 2008) and northeastern Kansas (Riley County, CNC), south to the state of Guanajuato on the Mexican Plateau (Ball and Shpeley 1992a: 60). The record from southern Wisconsin (Rauterberg 1885: 12) is probably in error.
Worldwide, with about 980 species (Lorenz 2005: 328-342) arrayed in two subtribes:
Worldwide, with about 900 species (Lorenz 2005: 328-341). The vast majority of species (about 81 % of the world fauna) are found in the Oriental and Afrotropical Regions.
Worldwide, with about 855 species (Lorenz 2005: 328-341) arrayed in 62 subgenera. The genus is mainly represented in the Afrotropical and Oriental Regions with about 650 species (approximately 76% of the world fauna). The North American fauna has 52 species (6% of the world fauna) in the boreal, temperate, and subtropical regions.
Bell (1960) revised the North American species. Since the publication of this work, one Neotropical species was discovered in southern Texas (
One species along the Coastal Plain of eastern North America.
This species is found from southeastern Georgia (Fattig 1949: 44) to southern Florida (Bell 1960: 102), west to southeastern Texas (Aransas County, USNM); also recorded from South
Seven species in the Nearctic (two species) and Neotropical (five species) Regions.
To date this species has been recorded only from southeastern Arizona (Bell 1960: 103).
This species ranges from Maine to southern Alberta, south to southeastern Arizona, central Texas, and central Florida (Bell 1960: 104-105).
Three species in eastern North America.
The generic name
This species ranges from Virginia (Robert L. Davidson pers. comm. 1992) to northeastern Iowa (Purrington et al. 2002: 201), south to eastern Texas (Sabine County, CMNH), southeastern Louisiana (Colby 2002: 37), and northern Florida (Bell 1960: 106).
This species ranges from Nova Scotia (Lindroth 1969a: 975) to eastern South Dakota (Kirk and Balsbaugh 1975: 36; French et al. 2004: 557), south to central Oklahoma and central Florida (Bell 1960: 107). Old specimens labeled “Tex” are also known (Bell 1960: 107).
This species ranges from southern Quebec (Saint-Clet, CNC) to southeastern Minnesota (Gandhi et al. 2005: 929), south to southern Texas and central Florida (Bell 1960: 108). At least one specimen labeled “Mass” is known (Bell 1960: 108).
About 140 species in all zoogeographical regions except the Australian Region. The Nearctic Region is represented by 14 species.
This species ranges from Massachusetts (Bell 1960: 120) to east-central Iowa (Wickham 1888: 82), including southernmost Ontario (Lindroth 1969a: 981), south to southeastern Louisiana (Allen 1965: 75) and central Florida (Bell 1960: 120). The record from eastern Kansas (Popenoe 1877: 23) needs confirmation.
This species ranges from New Jersey (Bell 1960: 121) to east-central South Carolina (Ciegler 2000: 80) and southwestern Alabama (Löding 1945: 23; Bell 1960: 121). The record from “Mississippi” (Bousquet and Larochelle 1993: 204) needs confirmation.
This species ranges from southern Texas (Johnson 1978: 68) to Veracruz (Davidson 1980: 88) along the east coast of Mexico.
This species ranges from “Massachusetts” to central North Dakota (Bell 1960: 122), north to southeastern Manitoba (Ryan and Holliday 2006: 414), south to northeastern New Mexico, central Texas, northwestern Georgia (Fattig 1949: 44) along the Appalachians, and Long Island, New York, along the east coast (Bell 1960: 122).
This rarely collected species is known from Arkansas, southern Missouri (Bell 1960: 121), and eastern Oklahoma (Latimer County, UASM). Specimens labeled «S[outh] Ill[inois]» are known (Bell 1960: 121).
This species is known from southeastern Arizona to central Texas (Bell 1960: 118), south to Durango and San Luis Potosí in Mexico (Davidson 1980: 133).
This species is known from Sonora (Davidson 1980: 129), southern Arizona, southern California (Bell 1960: 117), and the Baja California Peninsula (Horn 1894: 311).
This species is known from southern Alabama, southern Georgia, and throughout Florida (Bell 1960: 118). A specimen labeled “N. Car.” is known (Bell 1960: 118).
This species was known for a long time under the name
This species is known from the Baja California Peninsula and from southern Arizona south, through central and western Mexico, to Central America (Robert L. Davidson pers. comm. 2007).
Horn (1876d: 275) listed this name as a junior synonym of
This species ranges from New Jersey (Bell 1960: 131) to eastern South Dakota (Kirk and Balsbaugh 1975: 36), including southernmost Ontario (Lindroth 1969a: 982), south to northeastern New Mexico (Union County, CNC), southeastern Texas, and southern Florida (Bell 1960: 131).
This species is found in southeastern United States from South Carolina (Kirk 1970: 16; Ciegler 2000: 80) to northern Arkansas, south to southeastern Texas and southeastern Georgia (Bell 1960: 130). The records from Illinois (Leng and Beutenmüller 1893: 143; Blatchley 1910: 166), “Iowa” (Jaques and Redlinger 1946: 296), Kansas (Popenoe 1877: 23; Snow 1903: 193), “Oklahoma” (Arnold 2008), and Missouri (Summers 1873: 145) need confirmation; that from southeastern Pennsylvania (Rathvon 1869: 525) must be in error.
This species ranges from southern Maine (Majka et al. 2011: 47) to southeastern Nebraska (Nemaha County, Foster F. Purrington pers. comm. 2012), including southernmost Ontario (Bousquet 1987a: 132), south to Durango and Nuevo León (Davidson 1980: 92) in Mexico and southern Florida (Bell 1960: 129), west along southern United States to southeastern Arizona (Dajoz 2004: 116). The record from Tabasco in Mexico (Davidson 1980: 92) is suspect; that from southern Wisconsin (Rauterberg 1885: 18) needs confirmation.
This species ranges from southeastern Arizona to southeastern Texas, south to Oaxaca in Mexico (Bell 1960: 128); also recorded from San Bernardino County in southeastern California (Dajoz 2007: 20).
This transamerican species ranges from Newfoundland (Lindroth 1955a: 135) to southwestern British Columbia (Lindroth 1969a: 983), south to southern California (Bell 1960: 125, 127), Chihuahua (Davidson 1980: 84) and northern Sonora (Bates 1884: 267, as
Bell (1960: 123-128) recognized four subspecies within
Nineteen species can be assigned to this subgenus: five in North America (Bell 1960: 109, as
Liu et al. (2011: 22) argued that it is premature to place any of the Western Hemisphere species of
This species ranges from New Brunswick (Restigouche and Queens Counties, CNC) to southern Manitoba (Lindroth 1969a: 981), south to northwestern Arkansas (Bell 1960: 112), northwestern Mississippi (Coahoma County, Drew A. Hildebrandt pers. comm. 2008), and North Carolina (Robert L. Davidson pers. comm. 1992). The record from southwestern Colorado (Elias 1987: 634) needs confirmation.
This subspecies is found in the drainage basin of the Colorado River in southwestern Colorado, southern Utah, Arizona, and southeastern California (Bell 1960: 114).
This subspecies ranges from central Colorado to southernmost New Mexico and central Texas (Bell 1960: 115), including northwestern Oklahoma (Cimarron County, CMNH, CNC).
This subspecies is known from southwestern Oregon (Hatch 1953: 163, as
This species ranges from Massachusetts to “Colorado” (Horn 1876d: 267), including southern Michigan and “Minnesota” (Gandhi et al. 2005: 929), south at least to central Texas and southern Florida (Bell 1960: 110-111).
This species ranges from central Missouri (Morgan County, CMNH) to Arizona (Bell 1960: 110), south to Nicaragua (Bates 1882a: 45); also found along the Baja California Peninsula (Davidson 1980: 69).
This species inhabits the Great Plains and the Great Central Plains from southern Alberta to southwestern Pennsylvania (Allegheny and Westmoreland Counties, Robert L. Davidson pers. comm. 2008), south to northern Mississippi (Drew A. Hildebrandt pers. comm. 2007), east-central Louisiana (West Feliciana Parish, Igor M. Sokolov pers. comm. 2009), central Texas, and central New Mexico (Bell 1960: 111). The records from southwestern Ohio (Dury 1906: 257) and “Utah” (Bousquet and Larochelle 1993: 206) need confirmation. The record from “Georgia” (J.E. LeConte 1849: 26) is probably in error; that from Beaver Islands in Michigan (Hatch 1925: 553) was based on misidentified specimens of
About 80 species in all zoogeographical regions except the Australian Region. The subgenus is best represented in the Northern Hemisphere with 58 species (about 73.5% of the world fauna) of which 17 are found in North America.
This species ranges from southern Georgia (Fattig 1949: 43) and throughout Florida west along the Gulf Coast to southeastern Texas, north to north-central Oklahoma (Alfalfa County, CMNH) and central Arkansas (Pulaski County, Robert L. Davidson pers. comm. 2012), south at least to Veracruz along the Gulf Coast (Bell 1960: 146, as
This species has been known for a long time under the name
This species is found from central California (Bell 1960: 144) to western Texas (Jeff Davis, Ken Karns pers. comm. 2009), south to northern Sonora (Bates 1884: 267).
This species is known only from a few specimens collected in the lower Sacramento River Valley of northwestern California (Bell 1960: 145).
This species ranges from New Brunswick (Webster and Bousquet 2008: 21) to North Dakota (Tinerella 2003: 635), north to southern Manitoba (Lindroth 1969a: 984), south to eastern Texas (Houston County, CNC; Riley 2011) and southern Florida (Bell 1960: 137). The records from “Wyoming” and “Colorado” (Bousquet and Larochelle 1993: 207) are likely in error.
The name
This species ranges from south-central New Hampshire (Merrimack County, Ross T. Bell pers. comm. 2008) to eastern South Dakota (Kirk and Balsbaugh 1975: 35), including southern Ontario (Lindroth 1969a: 987), south to southern Texas (Bell 1960: 143) and southwestern South Carolina (Hampton County, CMNH). The records from “Utah” (Bousquet and Larochelle 1993: 207) and New Mexico (Fall and Cockerell 1907: 160) need confirmation.
This species ranges from Massachusetts (Easton 1909: 37) to northeastern South Dakota (Kirk and Balsbaugh 1975: 36), including southern Ontario (Lindroth 1969a: 984), south to central Texas and southern Florida (Bell 1960: 140). The record from “New Hampshire” (Bousquet and Larochelle 1993: 208) needs confirmation.
This species occurs from northwestern Georgia and southwestern South Carolina (Hampton County, CMNH) to central Florida, west to eastern Texas, north to central Oklahoma (Grady County, CMNH) and east-central Arkansas (Bell 1960: 141).
This subspecies ranges from Newfoundland (Lindroth 1955a: 134) to northwestern Montana (Russell 1968: 66), south to central New Mexico (Fall and Cockerell 1907: 160), central Texas, northwestern Georgia, and northern South Carolina (Bell 1960: 142).
Bell (1960: 142) noted the presence of intergrade specimens between the two forms of
This subspecies ranges from southern British Columbia, including Vancouver Island (Lindroth 1969a: 986), to southern California (Bell 1960: 142) and Guatemala (Bates 1882a: 44).
This species is known only from a few localities in central and southern Texas (Bell 1960: 149). The record from Georgia (Fattig 1949: 43) is likely in error (Bell 1960: 149).
This species is known from northern Georgia (Fattig 1949: 43) to southern Florida (Peck and Thomas 1998: 20) and from Inagua in the Bahamas (Darlington 1953: 8).
This species is found from southern Saskatchewan and southern Alberta (Lindroth 1969a: 988) south to southern Arizona and southern Texas (Bell 1960: 148).
This subspecies ranges from northern Utah to southern Arizona, east to western Texas (Bell 1960: 147).
This subspecies ranges from Newfoundland (Lindroth 1955a: 134) to Vancouver Island (Lindroth 1969a: 987), south at least to southwestern Oregon, southern Colorado along the Rocky Mountains (Bell 1960: 147), southeastern Louisiana (Saint Tammany Parish, Igor M. Sokolov pers. comm. 2009; Summers 1874a: 80), and southern Florida (Peck and Thomas 1998: 20). The record from southwestern California (Moore 1937: 12) is probably in error.
This species is endemic to the Florida Peninsula (Bell 1960: 165; Peck and Thomas 1998: 20).
This species is known from southern Mississippi (Hancock County, Drew A. Hildebrandt pers. comm. 2008), southern Texas, and northern Mexico (Bell 1960: 151).
This species ranges from southeastern Nebraska (Lancaster County, Foster F. Purrington pers. comm. 2010) and central Missouri (Boone County, CMNH) south to Mississippi (CMNH), southwestern Louisiana (Hine 1906: 77), and southern Texas (Bell 1960: 151). The record from eastern South Dakota (Kirk and Balsbaugh 1975: 36) needs confirmation.
This species ranges from central California (Bell 1960: 150) south to “Nicaragua” (Bates 1882a: 44) and the Baja California Peninsula (Horn 1894: 311), including western New Mexico (Catron County, UASM; Snow 1885: 67; Fall and Cockerell 1907: 160) and western Texas (Jeff Davis County, UASM). The record from “Utah” (Bousquet and Larochelle 1993: 208) needs confirmation.
The range of this species extends from southeastern Oregon (Harney County, James R. LaBonte pers. comm. 1992) and west-central Nevada (Bell 1960: 150) south to the Baja California Peninsula (Horn 1894: 311) and at least Durango in Mexico (Ball and Shpeley 1992a: 61). The specimens labeled from Texas (Bell 1960: 150) are probably mislabeled or represent strays.
Three species in southwestern North America (one species) and Mexico (three species).
This species ranges from southern California to southwestern New Mexico, south to Oaxaca in Mexico (Bell 1960: 116).
The name
Two species in the boreal and temperate regions of North America.
This species is known only from the type locality in north-central California (Bell 1960: 139).
Bell (1960: 139) stated that the holotype and still only known specimen may simply be an aberrant individual of
This species ranges from Newfoundland (Lindroth 1955a: 135) to central British Columbia, north to Fort Smith in southern Northwest Territories (Lindroth 1969a: 989), south to northeastern Washington (Hatch 1953: 163), “New Mexico” (Robert L. Davidson pers. comm. 1992), northern Texas (Cooke County, CNC), and northeastern Georgia (Bell 1960: 138). The record from “Oregon” (Wickham 1896c: 135) needs confirmation.
Nine species in the Nearctic (five species), Neotropical (one Mexican species also found in Arizona), and Palaearctic (four species) Regions.
This species ranges from Newfoundland (Larson and Langor 1982: 594) to central Alaska (Lindroth 1969a: 993), south to northeastern Washington (Hatch 1953: 162), central Colorado along the Rocky Mountains, western South Dakota (Meade County, Foster F. Purrington pers. comm. 2010), and eastern Minnesota (Bell 1960: 134). The species is unknown east of the Mississippi south of the Canadian border. Fossil remnants of this species, dated between about 16,700 and 25,200 years B.P., have been unearthed in southeastern Iowa (Baker et al. 1986: 96) and southeastern Illinois (Schwert 1992: 78).
This species is known from a few specimens simply labeled from Arizona and from Mexico as far south as Oaxaca and central Veracruz (Bell 1960: 132; Davidson 1980: 31).
This species ranges from southeastern Alberta to Vancouver Island (Lindroth 1969a: 990-991), south at least to central California and northern Utah (Bell 1960: 133). One old specimen labeled from “Arizona” is also known (Bell 1960: 133).
This species is restricted to the Pacific Coast ranging from southern British Columbia, including Vancouver Island (Lindroth 1969a: 993), south through the Coast Range to northern California (Bell 1960: 135). The record from Inyo County, California (Dajoz 2007: 19) needs confirmation. Fossil remnants from a Plio-Pleistocene sequence have been unearthed in northwestern Greenland (Böcher 1995: 25).
This species ranges from Newfoundland (Lindroth 1955a: 133) to south-central British Columbia (Lindroth 1969a: 992), south to northeastern Washington (Hatch 1953: 162), “Wyoming” (Robert L. Davidson pers. comm. 1992), east-central Texas, southern Florida, and Cuba (Bell 1960: 133-134). The record from “Yukon Territory” (Ball and Currie 1997: 453) could not be confirmed.
The name
One North American species.
In my opinion,
This species is found from southern Quebec (Lindroth 1969a: 994) to southeastern British Columbia (CNC), north to Fort Smith in southern Northwest Territories (Bousquet 1987a: 132), south to Oregon (Bell 1960: 136), New Mexico (Perrault 1970: 56), Kansas (Popenoe 1878: 78; Bell 1960: 136), and “New Jersey” (Smith 1890: 92; Smith 1910: 213).
Worldwide, with about 13,200 species placed in the following 32 tribes:
Worldwide, with about 235 species arrayed in four subtribes:
Forty-five species arrayed in two genera, both represented in the Nearctic Region.
Twenty-nine species in the boreal, temperate, and tropical areas of the Nearctic (nine species), Oriental (12 species), Palaearctic (nine species), and Afrotropical (two species) Regions. The species are placed in three subgenera:
Ball (1959) revised the species of this genus. Since this publication, one new North American species has been described by Will (1998) and a few nomenclatural changes affecting the Nearctic species have been introduced by Lindroth (1968-1969a): Ball’s
Eighteen species in the Nearctic (nine species), Oriental (three species), Palaearctic (five species), and Afrotropical (two species, one shared with northern Africa and Spain) Regions. Several species listed in this subgenus by Lorenz (2005: 342-343) belong to the subgenus
This species is found from New Brunswick (Webster and Bousquet 2008: 20) to east-central South Dakota (Kirk 1971: 239), south to “Georgia” (LeConte 1844: 51).
This species is known from a few localities in the Florida Panhandle and along the Mississippi Basin in northwestern Mississippi, western Tennessee, and southern Illinois [see Will 1998: Fig. 10].
This species ranges from southern Quebec (Larochelle 1975: 80) to west-central South Dakota (Kirk and Balsbaugh 1975: 34), south to central New Mexico (Ball 1959: 71, as
This subspecies ranges from “Rhode Island” to east-central South Dakota (Ball 1959: 78), south to east-central Kansas (Snow 1880: 78; Ball 1959: 78), “Missouri” (Ball 1960b: 78), and New Jersey (Smith 1890: 82; Smith 1910: 206). The records from Louisiana (Summers 1874a: 80; Casey 1913: 148) and Alabama (Löding 1945: 18) refer to subspecies
This subspecies ranges along the Coastal Plain from eastern Virginia (Hoffman et al. 2006: 24) to southern Florida (Peck and Thomas 1998: 21), west to southeastern Texas (Ball 1959: 80); also recorded from Cuba (Ball 1959: 80).
As far as known, this species is endemic to the southern half of the Florida Peninsula (Ball 1959: 73).
This species occurs from Cape Breton Island (Bousquet 1987d: 107) to south-central British Columbia, north to central Northwest Territories along the Mackenzie River (Lindroth 1969a: 948), south to eastern Washington, “Nevada” (Ball 1959: 82), the Sierra Blanca in south-central New Mexico (Fall and Cockerell 1907: 159), northwestern Texas (Oldham County, Foster F. Purrington pers. comm. 2010), “Missouri” (Ball 1959: 82), northwestern Tennessee (Stewart County, Foster F. Purrington pers. comm. 2010), and western Virginia (Hoffman et al. 2006: 24). The record from southern Louisiana (Summers 1874a: 80) is probably in error; that from “Arkansas” (Bousquet and Larochelle 1993: 209) needs confirmation.
This species ranges from southeastern Manitoba to the Okanagan Valley in southern British Columbia (Lindroth 1968: 943), south to northwestern Oregon (Hatch 1951: 119), “Utah” (Ball 1959: 64), northwestern Wyoming (Sikes 1994), and “North Dakota” (Donald P. Schwert pers. comm. 1989).
This species ranges from central Nova Scotia (Majka et al. 2007: 10) to the Okanagan Valley in British Columbia, north to Fort Smith in southern Northwest Territories (Lindroth 1968: 941), south to Inyo County in California (Dajoz 2007: 19), southeastern Arizona (Dajoz 2007: 21, as
This species is known from Alberta, as far north as Edmonton, southern Manitoba (Lindroth 1969a: 946), Minnesota (Tinerella and Rider 2001: 321; Kamal J.K. Gandhi pers. comm. 2008), Wisconsin (Messer 2010: 39), “Illinois” (Ball 1959: 83), and southeastern Nebraska (Hall County, UASM).
Sixteen species (24 species-group taxa) restricted to the temperate and tropical zones of North America and Mexico (south to the vicinity of Mexico City). Three subgenera are recognized:
Ball (1959) revised the species of this genus. Since the publication of this work, Ball (1992a) described one new Mexican species (
Seven North American species arrayed in three species groups.
This species ranges from Maine (Foss 2001: 14) to eastern South Dakota (Kirk and Balsbaugh 1975: 35), including southern Ontario (Lindroth 1969a: 950), south to east-central Texas (Riley 2011) and southern Florida, including the Keys (Peck and Thomas 1998: 21). The record from Montreal, Quebec (Perrault 1977: 153) needs confirmation.
This subspecies ranges from “Maine” (Larochelle and Larivière 1990a: 32) to west-central New York, south to northern Florida (Ball 1959: 131-132), west to eastern Texas (Sabine County, Brian Raber pers. comm. 2010).
Ball (1959: 132) reported the presence of intergrade populations of this and the
This subspecies ranges from western Pennsylvania to southeastern Iowa (Ball 1959: 134) and west-central Illinois (Willand et al. 2011: 273 as
This subspecies ranges from eastern Minnesota (Gandhi et al. 2005: 929) south to eastern Texas (Sabine County, Brian Raber pers. comm. 2010) and southern Mississippi, east to northwestern Georgia (Ball 1959: 123).
This subspecies ranges from eastern Pennsylvania to west-central Illinois (McCravy and Willand 2008: 157), south to Tennessee (Ball 1959: 122) and South Carolina (Kirk 1969: 14; Kirk 1970: 15; Ciegler 2000: 84). The records from New York (Notman 1928: 229, as
Ball (1959: 122) reported the presence of intergrade populations between this and the
This subspecies is found from southernmost Ontario and southeastern Michigan south to southern Indiana and central Maryland (Ball 1959: 143-144).
This subspecies is found in the southern part of the Great Plains from northeastern Kansas to central Iowa, south to central Arkansas and eastern Oklahoma (Ball 1959: 142) including southwestern Illinois (Union County, CNC). The record from western Pennsylvania by LeConte (1859d: 53) is probably in error.
Ball (1959: 142) reported the presence of intergrade populations between all three subspecies of
This subspecies ranges from western Ontario to Saskatoon in Saskatchewan (Lindroth 1969a: 953), south to southern New Mexico (Ball 1959: 145), southeastern Nebraska (Lancaster County, Foster F. Purrington pers. comm. 2009), and southern Illinois (Ball 1959: 145).
This species is known from northern New Jersey (Smith 1910: 207) to southeastern Iowa, south to southeastern Texas (Ball 1959: 113) and the Florida Panhandle (Peck and Thomas 1998: 21).
This species ranges from southern Quebec (Larochelle 1975: 80) to northern Iowa, south to northern Alabama (Ball 1959: 111) and South Carolina (Ciegler 2000: 84). The record from “Florida” (Horn 1880c: 52) needs confirmation.
This species ranges from southern Quebec (Larochelle 1975: 80) to southwestern Indiana, south to northern Alabama (Ball 1959: 115), northern Georgia (Fattig 1949: 31), and northwestern South Carolina (Ciegler 2000: 85). The record from South Dakota (Kirk and Balsbaugh 1975: 35) is likely in error.
Six North American species, one of them extending into northern Mexico, arrayed in three species groups.
This species is known from eastern Texas south to southern Tamaulipas and Durango (Ball 1959: 167) in Mexico. Two specimens are known from northern Florida and “Virginia” (Ball 1959: 167).
This species is known from southeastern coastal Georgia south to southern Florida (Ball 1959: 164).
This species inhabits the southern part of the Great Plains from “Kansas” to southeastern Missouri (Ball 1959: 162), south to northwestern Mississippi (Coahoma County, Drew A. Hildebrandt pers. comm. 2008), “Louisiana” (LeConte 1853c: 389), and southern Texas (Ball 1959: 162). The records from Alabama (Löding 1945: 18), the Florida Panhandle (Harris and Whitcomb 1974: 99), Georgia (Fattig 1949: 31), and North Carolina (Brimley 1938: 122) need confirmation.
This species is known only from a few localities in the southern third of Florida, including the Keys (Ball 1959: 165).
This form has been considered as a subspecies of
The range of this subspecies extends from Massachusetts to western Wisconsin (Ball 1959: 156), south to southeastern Texas (Cameron County, Brian Raber pers. comm. 2010), southern Louisiana (Summers 1874a: 80; Allen 1965: 73), southwestern Alabama (Ball 1959: 156), and central Georgia (Fattig 1949: 30).
The range of this subspecies extends from “Minnesota” (Ball 1959: 158) to southwestern North Dakota (Tinerella 2003: 636), south to southeastern Arizona, southeastern Texas (Ball 1959: 158), and southeastern Louisiana (Allen 1965: 73), including west-central Illinois (Willand et al. 2011: 273).
This species has been recorded from “Georgia” (LeConte 1847: 422), the Florida Panhandle and Peninsula including the Keys (Peck and Thomas 1998: 21), and southwestern Alabama (Löding 1945: 18; Ball 1959: 159).
Following Lindroth (1969a: 954), I believe this taxon should be regarded as a valid species, not as a subspecies of
Five species in North America (three species) and Mexico (four species, two of them endemic).
This species is confined to the Huachuca and Chiricahua Mountains of southeastern Arizona [see Ball 1992a: Fig. 17].
This subspecies ranges from southeastern Alberta to southwestern South Dakota, south to southeastern Texas, Chihuahua in Mexico, and southeastern Arizona [see Ball 1992a: Fig. 18].
The subspecies
This species is known only from the Chiricahua Mountains in southern Arizona and the Sierra Madre Occidental in Chihuahua and Sonora [see Ball 1992a: Fig. 17].
Worldwide, with about 100 species in the Nearctic (14 species of
Several specimens of
Forty-eight species in the boreal, temperate, and tropical areas in the Nearctic (14 species), Neotropical (five species in Mexico, the West Indies, and Peru), Australian (two species), Oriental (one species), Palaearctic (27 species), and Afrotropical (three species) Regions. The species are arrayed in three subgenera:
Ball (1959) revised the North American species. No new North American species have been described since. Lindroth’s (1969a) key encompasses all North American species but
Twenty-four species in North America (eight species, one of them extending into the Bahamas), Mexico (three species, one of them endemic), Peru (one species), and the Palaearctic Region (14 species).
Erwin and Ball (2011: 407-409) published a key to the Western Hemisphere species.
This rarely collected species ranges from “Illinois” to eastern Kansas (Ball 1959: 206), south to Coahuila and Tamaulipas (Ball 1992a: 375) in northern Mexico, southeastern Louisiana (East Baton Rouge Parish, Igor M. Sokolov pers. comm. 2009), and western Alabama (Tuscaloosa County, UASM). The records from southwestern New Jersey (Smith 1910: 207), “Florida” (Leng 1915: 580), “Delaware” (Houghton 1905: 211), and “Colorado” (Csiki 1931: 903) need confirmation.
This subspecies ranges from south-central New York (Ball 1959: 213) to southeastern Nebraska (Lancaster County, UASM), including southern Wisconsin (Rauterberg 1885: 17; Purrington et al. 2002: 201), south to southern Texas (Gonzales County, CMNH; Brazoria County, Robert L. Davidson pers. comm. 2008) and southern Florida (Peck and Thomas 1998: 21); also recorded from the Bahamas (Darlington 1953: 8).
Ball (1959) recognized three subspecies within this taxon, two of them occurring north of Mexico. Because the differences between the two forms are largely based on color and that intermediate specimens are known from Louisiana (Ball 1959: 210), I prefer not to recognize the color forms as distinct subspecies.
This rarely collected species ranges from southwestern New Jersey (Smith 1910: 207) to central Kansas (Knaus 1898: 19), south to east-central Texas (Riley 2011), central Louisiana (Concordia Parish, CNC), and northern Florida (Peck and Thomas 1998: 21).
This species is found from southwestern New Brunswick (Webster and Bousquet 2008: 20) to eastern South Dakota (Kirk and Balsbaugh 1975: 35; French et al. 2004: 557), south to east-central Texas (Brazos County, CNC; Riley 2011), northern Georgia (Ball 1959: 216-217), and northwestern South Carolina (Ciegler 2000: 85).
This species ranges from southwestern British Columbia, including Vancouver Island (Lindroth 1969a: 962), south to southern California (Ball 1959: 200-201). One old specimen labeled “Nev.” is known (Ball 1959: 200).
This species ranges from Cape Breton Island to southwestern British Columbia, north to the Great Slave Lake in Northwest Territories (Lindroth 1969a: 959), south to south-central California (Ball 1992a: 375), northeastern Kansas (Popenoe 1878: 78, as
This species is found from Cape Breton Island (Bousquet 1987a: 132) to south-central British Columbia (Jarrett and Scudder 2001: 380), north to central Northwest Territories (Ball 1992a: 375), south to northeastern New Mexico (Ball 1992a: 375) along the Rocky Mountains, the Guadalupe Mountains in western Texas (Ball 1992a: 375), the upper peninsula of Michigan (LeConte 1878c: 594), southern Ontario (Lindroth 1969a: 961), and southern Maine (Majka et al. 2011: 46). The record from the Pacific Northwest (Hatch 1953: 961) refers to
This rarely collected species is known from southern Ontario (Lindroth 1969a: 959), northern Ohio (Ashtabula County, Harry J. Lee, Jr. pers. comm. 2008), southern Michigan (Clinton County, CMNH), northern and southwestern Indiana (LeConte 1847: 418; Ball 1959: 205), western Illinois, “Tennessee” (Ball 1959: 205), southwestern Alabama (Lindroth 1954b: 153), and Connecticut (New London County, Foster F. Purrington pers. comm. 2009). Most old (pre-1950s) records of this species refer to
Twenty-two species in the Nearctic (six species), Neotropical (one species shared with North America), Australian (two species, one of them endemic to New Caledonia), Oriental (one species shared with the Australian Region), Palaearctic (11 species), and Afrotropical (three endemic species) Regions.
This transamerican species ranges from Cape Breton Island to Vancouver Island, south to “Oregon” (Lindroth 1969a: 964-965), northern Utah (Ball 1959: 225), east-central South Dakota (Kirk and Balsbaugh 1975: 35), northeastern Illinois (Purrington et al. 2002: 200), and the District of Columbia (Ball 1959: 225). The record from “Nebraska” (Bousquet and Larochelle 1993: 213) needs confirmation.
This species occurs from Nova Scotia (Lindroth 1954c: 305) to western Minnesota (Gandhi et al. 2005: 929, as
Casey (1920: 206-207), followed by Ball (1959: 227) and Lindroth (1969a: 966), stated that the species described by LeConte in 1844 as
This species ranges from southwestern Quebec (Bousquet 1987a: 132) to southeastern Minnesota (Ball 1959: 226), north to southern Manitoba (Lindroth 1969a: 967), south to southeastern Texas (Ball 1992a: 377) and northwestern Alabama (Colbert County, CMNH). One specimen labeled from Creston, British Columbia in UBC seen by Lindroth (1969a: 967) is probably mislabeled.
This species is known from Rhode Island (Sikes and Webster 2005: 315) to southern Wisconsin (Messer 2010: 39), including southernmost Ontario (Lindroth 1969a: 966, probably only as strays), south to the Rio Grande in southeastern Texas (Ball 1992a: 376), the Florida Keys, and the Greater Antilles (Ball 1992a: 376-377); it is also known from a single specimen, possibly a stray, collected in southwestern New Brunswick (Webster and Bousquet 2008: 21) and from Quintana Roo in the Yucatán Peninsula (Ball 1992a: 376). The record from “Quebec” (Bousquet and Larochelle 1993: 213) is in error.
In the
This species is known from three females collected in southern Oregon (Ball 1992a: 377) and north-central California (Motschulsky 1859a: 158).
This species ranges from New Brunswick (Webster and Bousquet 2008: 20) to southern Manitoba (Lindroth 1969a: 964; Roughley et al. 2010: 230), south to central Nebraska (Loup County, Foster F. Purrington pers. comm. 2010), northern Indiana, and New Jersey (Ball 1959: 219).
This species has been referred to the subgenus
Worldwide, with about 2,725 species arrayed in four subtribes:
Worldwide, with about 345 species arrayed in 30 genera (Lorenz 2005: 348-353). The North American fauna is represented by 48 species (about 14% of the world fauna).
Ball and Bousquet (2000: 90), following Noonan (1973: 388), recognized two genus-groups among the North American taxa, the Notiobii (an unavailable name; referred to as the “Notiobioid main branch” by Noonan) for the genus
About 95 species in the Nearctic (eight species), Neotropical (47 species), Australian (28 species), and Afrotropical (ten species) Regions arrayed in three subgenera:
Fifty-three species in the Nearctic (eight species, four of them endemic), Neotropical (21 species), and Australian (28 species) Regions.
Noonan (1973) revised the North and Middle American species and provided a key for their identification.
This species ranges from southeastern Arizona to southeastern Texas, south to southern Mexico; it is also known from southern Baja California [see Noonan 1973: Fig. 139].
This species is endemic to the Florida Peninsula (Shpeley 2001: 118).
This form has been listed as a synonym of
This species is known from Connecticut (Krinsky and Oliver 2001: 212), Long Island, New York (Noonan 1973: 307), the coast of southern North Carolina (Brunswick County, Ken Karns pers. comm. 2009), and from “Kansas” and northern Arkansas south to southern Mississippi and eastern Texas [see Noonan 1973: Fig. 134]. The records from New Mexico (Snow 1885: 67; Fall and Cockerell 1907: 162, as
This species ranges from the mountains in Arizona and northern New Mexico south to northern Panama; also known from the Laguna Mountains in southern Baja California [see Noonan (1973: Fig. 137)].
This species ranges from Maine and southern Quebec to southwestern South Dakota (Larsen and Purrington 2010: 571), south to eastern Texas and northeastern Georgia (Shpeley 2001: 119), including “Colorado” (van Emden 1953b: 544) [see Noonan 1973: Fig. 135]. The previous records from Florida (Noonan 1973) refers to
This species ranges from the San Francisco Bay area to southwestern Alabama, including central Missouri (Shockley and Cline 2004: 281), south to central Mexico and southern California [see Noonan 1973: Fig. 145]. The species is adventive on several islands of Hawaii (Liebherr 2009: 403).
This species ranges from southern Quebec (Larochelle 1975: 51, as
This widely distributed species ranges from western Newfoundland (CNC) to South Dakota (Kirk and Balsbaugh 1975: 32), south to Panama and southern Florida, west to southeastern Arizona [see Noonan 1973: Fig. 144]; also known from single specimens collected in southeastern Saskatchewan (Ronald R. Hooper pers. comm. 2007) and northern Colorado (Larimer County, William L. Krinsky pers. comm. 2008). The species is also recorded from Bermuda (Hilburn and Gordon 1989: 677).
One North American species in the temperate regions.
The species is included in Lindroth’s (1968: 864) monograph.
Lindroth (1968: 864) regarded this taxon as a subgenus of
This species ranges from Nova Scotia (Lindroth 1954c: 308) and Prince Edward Island (Majka et al. 2008: 132) to southeastern South Dakota (Kirk and Balsbaugh 1975: 31), south to southern Kansas (Snow 1903: 194), southern Arkansas (Kraim 1983: 229), and northwestern North Carolina (Watauga County, USNM). The records from “Colorado” (Snow 1877: 17; Wickham 1902: 243) need confirmation.
About 50 species in North America (33 species, one of them adventive), Mexico (seven species, two of them endemic), and the Palaearctic Region (17 species) arrayed in nine subgenera:
Twenty-two species in the Nearctic (14 species, of which two extends into the Baja California Peninsula and one is adventive) and Palaearctic (nine species, including the Himalayas) Regions.
Noonan (1996) revised all the species. One Palaearctic species,
This Palaearctic species is adventive in North America where it is known from the Skeena River valley in central British Columbia (Spence and Spence 1988: 158) to northeastern Oregon, east to west-central Montana (Hansen et al. 2009: 353) [see Noonan 1996: Fig. 230]; the species has been recorded also from Sacramento County in California (Clark 1999: 202). The specimen simply labeled from Nevada and the one labeled from Dickenson County in Iowa (Noonan 1996: 113) are likely mislabeled. The first inventoried specimen collected on this continent was found in 1911 at Portland, Oregon (Noonan 1996: 38). The species is also adventive in New Zealand since 1938 (Larochelle and Larivière 2005: 35).
This species ranges from northern Idaho and northern Washington south to northern Baja California Peninsula. A few specimens are known also from central Arizona and southern New Mexico [see Noonan 1996: Fig. 225].
This western species ranges from central Alberta to Vancouver Island, south to the Baja California Peninsula, southern Arizona, and northern Colorado [see Noonan 1996: Fig. 240]. Specimens labeled from “New Mexico,” Dallas (Texas), Jalisco in west-central Mexico, “Nebraska,” Lake County in Illinois, and “New Jersey” are known.
This eastern species ranges from New Jersey to northwestern Missouri, north to southernmost Ontario, south to east-central Texas and the Florida Panhandle [see Noonan 1996: Fig. 244]. Three specimens labeled from “Colorado,” Santa Cruz County in Arizona, and Lane County in Oregon are known.
This species occurs from central British Columbia south to the Mexican border in California and the eastern edge of the Rocky Mountains in northern Colorado [see Noonan 1996: Fig. 244]. Two old specimens labeled from “Arizona” and “Texas” are known.
This species is found from southern Quebec to northeastern South Dakota (Kirk and Balsbaugh 1975: 31), south to northern Colorado (Miller and Peairs 2008: 34), northeastern Texas, and central Florida [see Noonan 1996: Fig. 247]; the species has been collected at three sites in Washington which suggest that it has been introduced into the Pacific northwest.
This species was included in the subgenus
This species occurs from Nova Scotia to southwestern British Columbia, south to southern California, southern Arizona, the Texas Panhandle (Michels et al. 2010: 743), east-central Louisiana (West Feliciana Parish, Igor M. Sokolov pers. comm. 2009), central Mississippi, and west-central South Carolina (Ciegler 2000: 90) [see Noonan 1996: Fig. 248].
This species ranges from Long Island, New York, south to northern Florida and southeastern Mississippi [see Noonan 1996: Fig. 246]. One specimen labeled from “Massachusetts” is known.
This species is found from Newfoundland to western Ontario, south to northwestern Arkansas, southern Mississippi, and northern Georgia, west to eastern Kansas and Nebraska [see Noonan 1996: Fig. 245]. Seven specimens labeled from “Texas,” “South Dakota,” “California,” southwestern Saskatchewan, and southeastern Alberta are also known (Noonan 1996: 135-138). Several specimens were recently caught in the southern Similkameen Valley in British Columbia (Smith et al. 2004: 96) suggesting that the range of the species extends farther west or that the species was introduced in the area; a misidentification is also possible.
This species ranges from Maine and southern Quebec to southeastern Minnesota, south to northern Arkansas (Kraim 1983: 176), central Alabama, and northwestern South Carolina (Ciegler 2000: 89) [see Noonan 1996: Fig. 235]. The record from “Colorado” (Leng 1920: 72) is probably in error.
This species ranges from Massachusetts to southeastern Minnesota (Gandhi et al. 2005: 930), south to southern Missouri, “Mississippi” (Drew A. Hildebrandt pers. comm. 2007), and central South Carolina (Ciegler 2000: 90) [see Noonan 1996: Fig. 237]. There are two specimens from more western sites: one from western Nebraska, the other one from Utah. Whether these specimens are strays or represent elements of permanent populations remain to be seen. The record from South Dakota (Kirk and Balsbaugh 1975: 31) needs confirmation.
This species, known only from relatively few specimens, ranges from New Hampshire to eastern Michigan, south to northern Ohio (Ashtabula and Huron Counties, Harry J. Lee, Jr. pers. comm. 2008) and southern Pennsylvania (Somerset County, Robert L. Davidson pers. comm. 2012) [see Noonan 1996: Fig. 236]. One specimen collected in central Missouri (Boone County, CMNH) and identified by Gerald R. Noonan in 1994 is known (Robert L. Davidson pers. comm. 2012).
This species has a disjunct distribution. In the east, the species is found from Cape Breton Island to southeastern Manitoba south to southern Indiana, southern Pennsylvania, and New Jersey; in the west, the species ranges from southern British Columbia south to southwestern California, east to northern Idaho [see Noonan 1996: Fig. 238].
This species ranges from Nova Scotia to southern British Columbia, south to southern Washington, Colorado along the Rocky Mountains, Nebraska, and eastern Tennessee and western North Carolina along the Appalachian Mountains [see Noonan 1996: Fig. 239]. A few specimens simply labeled from California, Kansas, and Mississippi are known. The record from southwestern Alabama (Löding 1945: 25) is probably in error.
Ten species in North America (nine species) and Mexico (four species, one of them endemic,
Noonan (1973) revised all species.
This species ranges from extreme southern Ontario (Bousquet 1987a: 130) to eastern Nebraska, south to southeastern Texas and west-central Georgia [see Noonan 1973: Fig. 165]. Three specimens labeled from “Colorado,” Arizona (Huachuca Mountains), and California (Los Angeles County) are known. The record from New Jersey (Smith 1890: 98) needs confirmation.
This species ranges from northeastern Nebraska to eastern Virginia (Hoffman et al. 2006: 26), south to northern Mississippi (Lafayette and Panola Counties, Paul K. Lago pers. comm. 2009), northwestern Louisiana, and northeastern Texas (Noonan 1973: 361, Fig. 159). The records from “North Carolina” (Leng 1920: 72), South Carolina (Kirk 1969: 13; Kirk 1970: 14; Ciegler 2000: 90), Georgia (Fattig 1949: 50), “Florida” (Noonan 1973: 361), and southwestern Louisiana (Hine 1906: 77) need confirmation.
This species ranges from northern Kansas to northeastern Kentucky (Fayette County, Robert L. Davidson pers. comm. 2012) south to western Alabama and Nuevo León in northeastern Mexico [see Noonan 1973: Fig. 158]. The western limit of the species range is along the Rio Grande in western Texas, although it has been recorded from one locality in southwestern Arizona (Snow 1907: 142). The species is also known from south-central Wisconsin (Sauk County, Robert L. Davidson pers. comm. 2012). The records from South Carolina (Kirk 1969: 13; Kirk 1970: 14; Ciegler 2000: 90) need confirmation.
This species ranges from southeastern Louisiana to southeastern Arizona, south to northern Mexico [see Noonan 1973: Fig. 164].
This species ranges from Tulare County in east-central California to southwestern Texas, south to the Federal District area of Mexico [see Noonan 1973: Fig. 160].
This species ranges from Long Island, New York, to northeastern Oklahoma (Foster F. Purrington pers. comm. 2010), south to east-central Texas (Riley 2011) and southern Florida [see Noonan 1973: Fig. 166].
This eastern species ranges from southern Quebec to west-central Minnesota, north to southwestern Manitoba (Lindroth 1968: 846), south to southeastern Texas and the Florida Keys [see Noonan 1973: Fig. 161]. One specimen labeled from Anaheim, California is known. The records from “Arizona” and “North Dakota” (Bousquet and Larochelle 1993: 217) need confirmation.
This species ranges from Long Island, New York, to central South Dakota (Kirk and Balsbaugh 1975: 31), including southern Ontario, south to northeastern Texas, northeastern Georgia (House and All 1981: 195; Morrill 1992: 181), and west-central South Carolina (Ciegler 2000: 90) [see Noonan 1973: Fig. 163].
This species ranges from Prince Edward Island to Yellowstone National Park in Wyoming, north to southern Manitoba (Lindroth 1968: 845), south to northern Colorado (Miller and Peairs 2008: 34), eastern Texas, and northern Florida [see Noonan 1973: Fig. 162]. The records from “Arizona” (Leng 1920: 72) and New Mexico (Fall and Cockerell 1907: 162) need confirmation.
Seven species in North America (six species) and Mexico (
Noonan (2001) revised the species and provided a key for their identification.
This western species is found from southeastern British Columbia south to the Mexican border in California, southwestern Utah, and central Colorado [see Noonan 2001: Fig. 51].
This species ranges from southern Nova Scotia to eastern Montana, north to southern Manitoba (Lindroth 1968: 839) and west-central Saskatchewan, south to southeastern Texas and northern Florida [see Noonan 2001: Fig. 49]. The species is also, quite unexpectedly, found in western Oregon, some 1200 km from the western edge of the species main distribution area (Noonan 2001: 313).
This species ranges from southeastern Manitoba (CNC) to northwestern Washington, south to the Los Angeles area in California and, along the Rocky Mountains, to Arizona and northern New Mexico [see Noonan 2001: Fig. 44].
This species occurs from northern Idaho and eastern Washington, south to central California [see Noonan 2001: Fig. 49]. The records from New Mexico (LeConte, 1859c: 2; Fall and Cockerell 1907: 162) and “Arizona” (Leng 1920: 73) need confirmation. One specimen labeled “V[ancouver] I[sland]” is known (Lindroth 1968: 842, as
This species ranges from southwestern British Columbia to northwestern Wyoming (Park County, Ken Karns pers. comm. 2009), south to northeastern Nevada and the Mexican border in California [see Noonan 2001: Fig. 56].
This species ranges from the Sept-Iles region in easternmost Quebec to Vancouver Island, south to the Sierra Nevada in east-central California, southern Colorado in the Rocky Mountains, Oklahoma, southern Louisiana, and northern Florida; seemingly isolated in northwestern British Columbia [see Noonan 2001: Fig. 50].
One North American species in the temperate regions.
Lindroth (1968: 863-864) covered the species.
Lindroth (1968: 862) and Noonan (1973: 374) treated this taxon as a subgenus of
This species ranges from New Brunswick (Webster and Bousquet 2008: 19) to southern Saskatchewan (Roche Percee, CNC), south to east-central Texas (Riley 2011), northwestern Louisiana (Natchitoches Parish, Igor M. Sokolov pers. comm. 2009), northern Georgia (Fattig 1949: 52), and northwestern South Carolina (Ciegler 2000: 91).
Two North American species in the temperate regions.
Both species are included in Lindroth’s (1968: 857, 860) monograph.
This species is found west of the Rocky Mountains, from south-central British Columbia (Lindroth 1968: 857) south to southern California (Fall 1901a: 51; Moore 1937: 13).
This species ranges from “Massachusetts” (LeConte, 1847: 386, as
One eastern North American species in the temperate and subtropical regions.
The species is covered in Lindroth’s (1968: 860) monograph.
This species occurs from southeastern New York (Notman 1928: 248) to eastern South Dakota (Kirk and Balsbaugh 1975: 31), including southernmost Ontario (Bousquet 1987a: 131), south to east-central Texas (Riley 2011; Horn 1880d: 175) and southern Florida (Peck and Thomas 1998: 21).
One North American species in the temperate regions.
The species is covered in Lindroth’s (1968: 832-833) monograph.
This taxon has always been treated as a valid genus. In a phylogenetic analysis based on characters of the adult conducted by Noonan (1973: Fig. 241), this taxon nested within the genus
This widely distributed species ranges from Maine (Robert E. Nelson pers. comm. 1989) and southern Quebec (Larochelle 1975: 85) to southwestern Idaho and northwestern Nevada, north to the southern parts of the Prairie Provinces, south to northern Arizona (Johnson and Clark 1989: 443; Fig. 1), southern Texas (Johnson 1978: 67), and southern Georgia (Fattig 1949: 46).
Two North American species arrayed in two subgenera.
Lindroth (1968: 860-861) covered both species in his treatment of the genus
Lindroth (1968: 860) regarded this taxon as a subgenus of
One North American species in the temperate regions.
This species ranges from New Brunswick (Webster and Bousquet 2008: 19) to western South Dakota (Kirk and Balsbaugh 1975: 32), north to southern Saskatchewan (Ronald R. Hooper pers. comm. 1990), south to east-central Texas (Riley 2011), south-central Louisiana (Evangeline Parish, MCZ) and the Florida Panhandle (Liberty County, CNC).
One North American species in the temperate regions.
This species ranges from southern Quebec (Larochelle 1975: 49) to southeastern South Dakota (Kirk and Balsbaugh 1975: 32), south to east-central Texas (Riley 2011), northern Arkansas (Kraim 1983: 221), and west-central South Carolina (Kirk 1970: 14; Ciegler 2000: 92).
Five North American species, of which two extend into northern Baja California and Guadalupe Island.
Noonan (1968) revised the species and provided a key for their identification.
This species is known from east-central Washington (Casey 1924: 134, as
This subspecies is known from a few localities in southernmost California and northern Baja California [see Noonan 1968: Fig. 19].
This subspecies is found over most of California, south to the Los Angeles area and San Clemente Island [see Noonan 1968: Fig. 19].
This species is known from northeastern Oregon south to Los Angeles County in southern California [see Noonan 1968: Fig. 15].
This species ranges from Vancouver Island to northwestern Montana (Russell 1968: 67), south to northwestern Utah and the Mexican border in California [see Noonan 1968: Fig. 18]; also recorded from Guadalupe Island in the Pacific (Noonan 1968: 289).
This species is known only from a few localities in Kern and Tulare Counties in southern California [see Noonan 1968: Fig. 17].
New World, with about 90 species arrayed in eight genera:
In the phylogenetic analysis of the tribe
Twenty-eight species in the Nearctic (two species) and Neotropical (28 species) Regions arrayed in three subgenera:
Ten North and Middle American species.
Goulet (1974b) revised the species and provided a key for their identification.
The range of this species extends from southeastern Arizona and southwestern New Mexico south to Guatemala [see Goulet 1974b: Fig. 32].
This subspecies is known only from the holotype collected in southern Arizona.
The subspecies
Worldwide, with about 625 species (Lorenz 2005: 353-362) arrayed in approximately 35 genera. The Northern Hemisphere is represented by about 325 species (roughly 52% of the world fauna) and North America alone by 102 species (16.3%).
Ball and Bousquet (2000: 90) recognized two genus-groups among the North American taxa, the Polpochili for the genus
About 170 species (Lorenz 2005: 353-355, excluding
About 60 species in the Nearctic (15 species), Neotropical (ten species), Oriental (11 species), and Palaearctic (about 25 species) Regions.
Lindroth (1968: 905-907) treated all currently recognized valid species in his key except for
This species ranges from Vancouver Island (Lindroth 1968: 911) to western Montana (Russell 1968: 67), south to southern Colorado (Elias 1987: 634) and southern California (Fall 1901a: 50; Andrews et al. 1979: 28).
The range of this species extends from northern New Hampshire (Reeves et al. 1983: 459) to eastern South Dakota (Kirk and Balsbaugh 1975: 33), including southernmost Ontario (Lindroth 1968: 908), south to southern Texas (Johnson 1978: 67) and southern Florida (Peck and Thomas 1998: 22). The record from Nova Scotia (Lindroth 1954c: 309) is considered doubtful by Majka et al. (2007: 10).
This species ranges from southeastern California (Imperial County, MCZ, USNM) to westernmost Texas (Casey 1914: 279, as
This species ranges from northeastern Ohio (Purrington et al. 1999: 47) to southern Wisconsin (Rauterberg 1885: 21; Messer 2010: 40), north to Charity Island in the Michigan waters of Lake Huron (Andrews 1916: 79), south to southern Texas (Johnson 1978: 67), northern Georgia (Fattig 1949: 54), and northwestern South Carolina (Ciegler 2000: 94). The record from New Jersey (Smith 1910: 216) needs confirmation.
This species is known from northern California (Shasta County, USNM) to southern California (Fall 1901a: 50; Dajoz 2007: 20) and “Arizona” (Leng 1920: 74). The record from “Oregon” (Schaupp 1883a: 14) is probably in error (Hatch 1953: 184).
The range of this species extends from Newfoundland (Lindroth 1955a: 149) to Vancouver Island (Lindroth 1968: 910), south to the Sierra Nevada in California (Dajoz 2007: 16), northern Colorado (Haubold 1951: 706; Armin 1963: 122), and eastern Georgia (Fattig 1949: 54). The record from southern Louisiana (Summers 1874a: 81) needs confirmation.
This species occurs from southwestern New Brunswick (Webster and Bousquet 2008: 19) to eastern South Dakota (Kirk and Balsbaugh 1975: 33), south to northeastern Kansas (Popenoe 1878: 79) and south-central Maryland (Prince Georges County, USNM). The record from southern Colorado (Elias 1987: 634) needs confirmation.
This species ranges from southern Nova Scotia (Christopher G. Majka pers. comm. 2007) to southwestern Michigan (Allegan County, CMNH), south to southeastern Mississippi (Stone County, Drew A. Hildebrandt pers. comm. 2008), northern Georgia (Fattig 1949: 54), and eastern South Carolina (Ciegler 2000: 94).
This species is found from Vancouver Island (Lindroth 1968: 911) to northern Idaho (Hatch 1953: 183), south to “Arizona” (Casey 1914: 273, as
This species ranges from Vancouver Island (Lindroth 1968: 909) to northwestern Montana (Russell 1968: 67), south to central Utah (Casey 1914: 276) and southern California (Fall 1901a: 50; Moore 1937: 14). The species is adventive on Oahu, Hawaii (Liebherr 2009: 403).
This species is restricted to a small area from southern Quebec and the Ontario Peninsula (Bousquet 1987a: 131) south to central Pennsylvania (Clinton County, CMNH) and “Rhode Island” (Sikes 2003: 8).
This species ranges from Nova Scotia (Lindroth 1954c: 309) to southeastern Saskatchewan (Ronald R. Hooper pers. comm. 2007), south to southern Texas (Johnson 1978: 67) and southern Florida (Peck and Thomas 1998: 22), west along the southwest to southern California (Horn 1894: 312; Andrews et al. 1979: 28); also found in southern Baja California (Horn 1894: 312), the Bahamas (Darlington 1953: 10), Cuba (Darlington 1934: 112), Jamaica (Darlington 1941a: 14), Dominican Republic (Robert L. Davidson pers. comm. 2012) and Puerto Rico (Wolcott 1936: 191).
The range of this species extends from southern Quebec (Larochelle 1975: 255) to eastern Iowa (Johnson County, USNM), south to southeastern Texas (Aransas County, USNM) and southern Florida (Peck and Thomas 1998: 22); also recorded from Bermuda (Hilburn and Gordon 1989: 677). The records from central Colorado (Wickham 1902: 242) and Tabasco in Mexico (Bates 1891a: 246) are probably in error.
Besides the three original specimens, I assign to this species a series of specimens collected in southernmost Ontario (Rondeau Provincial Park, CNC).
This species is known from New Jersey (Smith 1910: 215; Lindroth 1968: 905) to the Florida Keys (Peck and Thomas 1998: 22), west along the Gulf Plain to southern Texas (Johnson 1978: 67), north to western Kansas (Scott County, CMNH).
Seven species in North America, of which two extend into northern Mexico, Bermuda, the Bahamas, and Cuba.
The subgenus has been reviewed by Heading (1964) but his work remains unpublished. Lindroth (1968: 916-920, as
This species is known from southern Oklahoma (Marshall County, CMNH) to eastern Coahuila in Mexico (UASM), east to southeastern Louisiana (Heading 1964: 34).
This species ranges from Newfoundland (Lindroth 1955a: 150) to Vancouver Island (Lindroth 1968: 919), south to southeastern California (Dajoz 2007: 20), southern Arizona, central Texas, northern Alabama, and east-central South Carolina (Ciegler 2000: 93) [see Heading 1964: Fig. 19]. The record from “Mississippi” (Bousquet and Larochelle 1993: 221) needs confirmation.
This species ranges from “New York” (LeConte 1847: 373) and New Jersey to southern Florida including the Keys (Peck and Thomas 1998: 22), west to northern Oklahoma (Alfalfa County, Robert L. Davidson pers. comm. 2008) and southeastern Texas [see Heading 1964: Fig. 18]; also recorded from the Bahamas and Cuba (Darlington 1953: 11). The records from southeastern Minnesota (Gandhi et al. 2005: 931) and southwestern Ohio (Blatchley 1910: 177) need confirmation.
The range of this species extends from western Maine (Kennebec and Somerset Counties, Ross T. Bell pers. comm. 2008) to eastern South Dakota (Ellsbury et al. 1998: 621), south to east-central Texas (Riley 2011) and the Florida Panhandle (Peck and Thomas 1998: 22). Also known from northern Idaho (Hatten et al. 2007: 359) and from a single, old (1926) specimen (possibly mislabeled) from western Oregon (Westcott et al. 2006: 9). The records from northern Colorado (Armin 1963: 115) and New Mexico (Fall and Cockerell 1907: 161,
This species is found from Prince Edward Island to southern Alberta (Lindroth 1968: 918), south to “Baja California” (Leng 1920: 74), southeastern California (Andrews et al. 1979: 28), southern Arizona (Heading 1964: Fig. 18), north-central Mexico (Heading 1964: 30), and central Florida (Peck and Thomas 1998: 22); also known from one specimen collected in northern Oregon (Westcott et al. 2006: 9) and reported as “common in light traps” in Bermuda (Hilburn and Gordon 1989: 677).
This species is found throughout California, in southeastern Arizona [see Heading 1964: Fig. 18], and in “Nevada” (LeConte 1869a: 374); one specimen labeled from south-central Washington is also known (Heading 1964: 32).
This species is known from central Washington to southwestern California [see Heading 1964: Fig. 19].
Seven North American species in the temperate regions.
There is no key for the identification of all species. Lindroth (1968: 921-924) covered four species, leaving
Adults and particularly larvae (personal observation) are structurally quite different from those of
This species ranges from Cape Breton Island (Lindroth 1954c: 309) to Vancouver Island (Lindroth 1968: 922), south to the Sierra Nevada in California (Dajoz 2007: 16), at least San Luis Potosí in Mexico (CNC), and central Florida (Peck and Thomas 1998: 22). The record from Guatemala (Bates 1882a: 71) needs confirmation.
This species ranges from east-central Ohio (Usis and MacLean 1998: 67) to southeastern Wyoming (Lavigne 1977: 47) and northeastern Colorado (Lavigne 1978: 102), including southernmost Ontario (Lindroth 1968: 924; Bousquet 1987a: 131), south to southeastern Texas (Casey 1914: 282) and northwestern South Carolina (Kirk 1970: 15). The record from “New York” (Bousquet and Larochelle 1993: 222) needs confirmation.
The range of this species extends from southern Quebec (Larochelle 1975: 109) to eastern Minnesota (Gandhi et al. 2005: 931), south to west-central Mississippi (Washington County, CMNH) and northwestern South Carolina (Kirk 1970: 15). The record from Colorado (Elias 1987: 634) is probably in error.
This species ranges from east-central Vermont to southeastern North Dakota, south to northeastern Kansas, Tennessee, and northeastern Virginia [see Bousquet and Messer 2010: Fig. 3].
This taxon has been listed in synonymy with
This species is known only from the type series collected in southwestern California.
I have studied both syntypes, including the male genitalia of one of them, and I have little doubt that they belong to a distinct species, though probably closely related to
This species is known from two localities in eastern Colorado and central New Mexico (Bousquet 1990: 203).
This species is known for sure only from southern California, as far north as the San Francisco Bay (Casey 1914: 280). The records from Colorado (Wickham 1902: 242; Elias 1987: 634) need confirmation; that from the Similkameen Valley in British Columbia (Smith et al. 2004: 96) was based on misidentified specimens of
About 130 species (Lorenz 2005: 356-358) arrayed in ten subgenera:
Noonan (1976: 21) also included
Eight species in North America (seven species) and Mexico (one species).
Fall (1905: 175, as
This species is known only from the type locality in west-central California.
This species is known from the Sierra Nevada and San Bernardino Mountains (Fall 1905: 176) and from San Diego County (Moore 1937: 14) in California.
This species is known only from the type locality in northwestern Nevada.
The range of this species extends from southwestern Washington (Hatch and Kincaid 1958: 6; Pacific County, MCZ) southwards to the Baja California Peninsula (Horn 1894: 312) and Guatemala (Ball and Shpeley 1992a: 58). The record from “British Columbia” (Horn 1894: 312) is likely in error (Lindroth 1968: 883).
This species is known so far only from southern California (Fall 1905: 176).
This species ranges from the Okanagan Valley in south-central British Columbia (Lindroth 1968: 883) to southern California (Fall 1905: 176; Dajoz 2007: 20).
This species is known only from the type locality in the Sierra Nevada.
About 45 species in the Nearctic (two species, one of them adventive), Neotropical (25 species), and Palaearctic (17 species) Regions.
The two species found in North America have been covered by Lindroth (1968: 883-884, as
This species is found along the Pacific Coast from northwestern Washington (Nelson 1988b: 56) to southwestern Oregon (Coos County, CNC).
This Palaearctic species is adventive in North America where it is known from the Vancouver area in southwestern British Columbia (Lindroth 1968: 883) south to Oregon (Westcott et al. 2006: 7; Lincoln County, Foster F. Purrington pers. comm. 2009). The first inventoried specimen collected on this continent was found in Vancouver in 1951 (Lindroth 1968: 883).
One North American species.
The species was covered in Lindroth’s (1968: 885) treatment of the Canadian
This species ranges from Newfoundland (Lindroth 1955a: 147) to eastern Alaska (Lindroth 1968: 885), south to northern Oregon (Hatch 1953: 182, as
Thirty-four species in North America (33 species), Mexico (three species, one endemic to Guadalupe Island), and the Bahamas (one species).
Casey (1914: 243-257) published a key to the species then known except
According to Kataev and Matalin (in Kryzhanovskij et al. 1995: 135), the sole syntype (a badly damage ♂) of
This species, as far as known, ranges from southeastern New Hampshire (Rockingham County, Ross T. Bell pers. comm. 2008) to northwestern Minnesota (Gandhi et al. 2005: 930), including southernmost Ontario (Lindroth 1968: 892), south to South Carolina (Charleston County, CNC).
This species ranges from Newfoundland (Lindroth 1955a: 147-148) to northeastern British Columbia (Lindroth 1968: 892), south to southwestern Oklahoma (Kondratieff et al. 2005: 171), southern Arkansas (Kraim 1983: 277), northeastern Georgia (Fattig 1949: 53), and southeastern South Carolina (Ciegler 2000: 96). The record from southwestern Alabama (Löding 1945: 26) needs confirmation; that from “Utah” (Bousquet and Larochelle 1993: 225) is probably in error.
This species ranges from New Brunswick (Lindroth 1968: 891) to North Dakota (Tinerella 2003: 636), north to southeastern Manitoba (Lindroth 1968: 891), south to east-central Arkansas (Kraim 1983: 280), northeastern Mississippi (Snodgrass and Cross 1983: 16), and southeastern South Carolina (Ciegler 2000: 96). The record from “New Mexico” (Fall and Cockerell 1907: 162) is probably in error.
This species is known only from the type locality.
The two syntypes are morphologically similar to adults of
This species ranges from Nova Scotia (Majka et al. 2007: 10) to Vancouver Island (Lindroth 1968: 888), south at least to central California (Clark 1999: 202; Dajoz 2007: 17), northern Sonora in Mexico (Bates 1884: 277), southern Texas (Johnson 1978: 67), southeastern Louisiana (East Baton Rouge Parish, Igor M. Sokolov pers. comm. 2009), Tennessee (Cumberland County, CNC), and southern West Virginia (McDowell County, Ken Karns pers. comm. 2009).
This species is known only from the type locality along the Pacific Coast of California.
This species is known from northwestern Georgia (Fattig 1949: 53) and “Texas” (Horn 1883a: 52). The record from Baja California (Horn 1894: 312) needs confirmation.
Fall (1905: 171) stated that
This species is known from a small area including southern Saskatchewan, southern Alberta (Lindroth 1968: 889), and southwestern North Dakota (Tinerella 2003: 636). The records from “Montana” and “Wyoming” (Bousquet and Larochelle 1993: 225) need confirmation.
The range of this western species extends from central Alberta to Vancouver Island (Lindroth 1968: 887), south to southern California and northern Sonora (LeConte 1858b: 60). The record from southern Texas (Johnson 1978: 67) needs confirmation.
This species ranges from Cape Breton Island (Lindroth 1954c: 308) to southeastern Wyoming (Lavigne 1977: 44), south to southern New Mexico (Fall and Cockerell 1907: 162), southern Texas (Wickham 1897: 113), southern Louisiana (Summers 1874a: 81; Allen 1965: 72, as
This species is known only from the holotype. Casey (1914: 258) assumed that the species was “possibly a native of California” because the holotype was sent to Chaudoir by Motschulsky. Although Motschulsky described several species from the west coast, he never set foot in western United States.
This species is found from Vancouver Island (Lindroth 1968: 894) to western Montana (Russell 1968: 68), south to west-central Nevada (Lyon County, CNC) and southern California (Fall 1901a: 50).
This species is known from “Texas” (Horn 1894: 312), Arizona (Griffith 1900: 566; Casey 1914: 249), southeastern California (Fall 1901a: 50), and the Baja California Peninsula (Horn 1894: 312).
This species ranges from southwestern Maine (Majka et al. 2011: 46) to eastern South Dakota (Kirk and Balsbaugh 1975: 33), including southern Ontario (Lindroth 1968: 893; Bousquet 1987a: 131), south to southeastern Texas (Fort Bend, Brian Raber pers. comm. 2010), northern Louisiana (Allen 1965: 72), southern Florida (Peck and Thomas 1998: 21), and the Bahamas (Darlington 1953: 10). The record from “Quebec” (Bousquet and Larochelle 1993: 226) was based on a misidentified specimen of
This species is known only from the type locality in northern California.
This species is known only from the type locality in southwestern California.
This species is known only from the type locality.
This species is known only from the type locality in southern Arizona.
This species is known only from the type locality in northwestern California.
This species is known only from the type locality in north-central California.
This species ranges from southern Montana (Hatch 1933a: 10) and South Dakota (Kirk and Balsbaugh 1975: 33) south to “Texas” (Casey 1914: 257) and southern Arkansas (Kraim 1983: 283). The record from Cuba (Darlington 1934: 110) is doubtful.
This species is known only from the type locality in northwestern California.
This species is known only from the type locality in westernmost Texas.
This species is known only from the type locality in western California.
This species is known only from the type locality in the Sierra Nevada.
This species is known only from the type locality in the Sierra Nevada.
This species is known only from the type locality in north-central Utah.
This species is known from the type series and San Diego County (Moore 1937: 14) in California.
This species is known only from the type locality in southwestern California.
This species is known for sure only from the type series collected in western California; it was also recorded from Humboldt County (Notman 1929b: 222).
This species is known for sure only from the type locality in east-central Texas. The record from South Dakota (Kirk and Balsbaugh 1975: 33) needs confirmation.
This species is known for sure only from the type series collected on the west coast in central California. Casey (1914: 254) reported it from Saint Helena, Sonoma County, California.
This species is known yet only from the type locality.
Two North American species.
Lindroth (1968: 894-898, as
This species extends from Newfoundland (Lindroth 1955a: 146) to southeastern Alaska, including Kodiak Island (Lindroth 1968: 896), south to mountains in southeastern California (Lindroth 1968: 896; San Bernardino County, CNC) and New Mexico (Fall and Cockerell 1907: 162), northwestern Indiana (Blatchley 1910: 193), and southeastern West Virginia (Greenbrier County, CNC). The record from “North Carolina” (Bousquet and Larochelle 1993: 226) needs confirmation.
This species ranges from Nova Scotia (Majka et al. 2007: 10) to central Minnesota (Gandhi et al. 2005: 930), north to southeastern Manitoba (Roughley et al. 2010: 230), south at least to east-central Missouri (Saint Louis, MCZ), southwestern Illinois (Saint Clair County, MCZ), and southeastern Pennsylvania (Dauphin County, MCZ). The single specimen from McMurray, Alberta (Lindroth 1968: 898) is possibly mislabeled or represents a stray.
Six North American species in the boreal and temperate regions.
Lindroth (1968: 898-903, as
The range of this species extends from New Brunswick (Lindroth 1968: 899) to eastern South Dakota (Kirk and Balsbaugh 1975: 32), north to southeastern Manitoba (Lindroth 1968: 899), south to “Texas” (Fall 1905: 172), Tennessee (Sevier County, CNC), and northeastern Georgia (Fattig 1949: 53).
This species is known from southwestern New Brunswick (Webster and Bousquet 2008: 19) to northern Minnesota (Gandhi et al. 2005: 930), including southern Quebec and the Ontario Peninsula (Lindroth 1968: 899), south to northeastern Kansas (Popenoe 1877: 24), Tennessee (Knox County, CNC), northern Georgia (Fattig 1949: 53), and eastern South Carolina (Ciegler 2000: 96). The records from Colorado (Wickham 1902: 242; Armin 1963: 119) are probably in error.
This species is restricted to the Pacific Coast from southern British Columbia, including Vancouver Island (Lindroth 1968: 903), south at least to the San Francisco Bay in California (Fall 1905: 173). The record from northern Colorado (Armin 1963: 119) is likely in error.
This species is yet known only from a few localities in Alberta.
This species ranges from southwestern New Brunswick (Webster and Bousquet 2008: 19) to central Minnesota (Sherburne County, CNC), including southern Quebec and the Ontario Peninsula (Lindroth 1968: 901), south to northeastern West Virginia (Randolph County, CMNH).
The range of this species extends from northeastern Newfoundland (Lindroth 1955a: 146, as
One North American species in the temperate regions.
The species was treated by Lindroth (1968: 877-878).
This species ranges from Massachusetts (Lindroth 1968: 878) to southeastern Wisconsin (Rauterberg 1885: 20), including southernmost Ontario (Bousquet 1987a: 131), south to southeastern Texas (Harris County, Brian Raber pers. comm. 2010), northern Louisiana (Allen 1965: 71, as
Forty-six species (Lorenz 2005: 358) in the Nearctic (two Holarctic species) and Palaearctic (46 species) Regions arrayed in five subgenera:
Lindroth (1968: 874-877) covered both species found in North America under the generic name
One Holarctic species.
This Holarctic subspecies ranges from eastern Siberia (Jaeger and Kataev 2003: 402) to Labrador (Lindroth 1954d: 370); isolated at high altitude in Colorado (Lindroth 1968: 877). Fossil remnants from the Early Wisconsinan have been unearthed in southern Ontario (Morgan and Morgan 1981: 1108).
Two other subspecies,
Thirty-one species in the Nearctic (one Holarctic species) and Palaearctic (31 species) Regions.
This circumpolar species ranges from Iceland to eastern Siberia (Jaeger and Kataev 2003: 403) and from Alaska (Lindroth 1968: 876) to Greenland (Böcher 1988: 14), south to southeastern Massachusetts (Cape Cod, CNC), central Iowa (Wickham 1911b: 8), central South Dakota (Kirk and Balsbaugh 1975: 32), the Sacramento Mountains in New Mexico (Fall and Cockerell 1907: 162), and the Baja California Peninsula (Horn 1894: 312). The records from southeastern New York (Notman 1928: 249) and “Pennsylvania” (Bousquet and Larochelle 1993: 227) need confirmation.
Worldwide, with about 125 species in the Nearctic (12 species), Neotropical (ten species), Australian (about ten species), Oriental (about 15 species), Palaearctic (about 50 species), and Afrotropical (30 species) Regions arrayed in eight subgenera:
Noonan (1976: 23) treated
Worldwide with about 75 species in the Nearctic (six species, one of them adventive), Neotropical (ten species), Australian (three species), Oriental (six species), Palaearctic (28 species), and Afrotropical (22 species) Regions.
Lindroth (1968: 929-934, as
This species ranges from Newfoundland (Lindroth 1955a: 149) to southern Saskatchewan (Lindroth 1968: 934), south to “South Dakota” (Lindroth 1955a: 149), northern Indiana (La Porte County, CMNH), and central Pennsylvania (Lycoming and Clinton Counties, CMNH). The record from “Delaware” (Bousquet and Larochelle 1993: 228) was based on a misidentified specimen of
The range of this species extends from Saint Pierre and Miquelon and southwestern Newfoundland (Lindroth 1955a: 148, as
This species ranges from southeastern New Hampshire (Rockingham County, Ross T. Bell pers. comm. 2008) to at least northern Illinois (Cook County, CMNH), south to northeastern Louisiana (Allen 1965: 72) and northern Florida (Alachua County, CNC).
This Palaearctic species is adventive in North America where it is known from the Quebec City area (Chantal 1971: 202) in the east and from south-central Saskatchewan (Ronald R. Hooper pers. comm. 2007) to Vancouver Island (Lindroth 1968: 930), including central Alberta (Pollock 1991a: 705), south to “Oregon” (Hatch 1953: 185) and northern Idaho (Hatten et al. 2007: 359) in the west. The first inventoried specimen collected on this continent was found in Seattle in 1931 (Hatch 1946: 77).
This species ranges from Nova Scotia (Lindroth 1968: 933) to northwestern Minnesota (Polk County, CMNH), north to southeastern Manitoba (Roughley et al. 2010: 230; CMNH), south at least to southwestern Pennsylvania (Allegheny County, CMNH) and Rhode Island (Casey 1914: 268). One old specimen simply labeled from Illinois is known (CMNH). The record from “Connecticut” (Bousquet and Larochelle 1993: 228) needs confirmation (see Krinsky and Oliver 2001: 4).
The range of this species extends from Nova Scotia (Lindroth 1968: 932) to northwestern Minnesota (Polk County, CMNH), south to west-central West Virginia (Mason County, CMNH) and Delaware (Lindroth 1968: 932). The record from eastern South Dakota (Kirk and Balsbaugh 1975: 33) needs confirmation.
Four North American species, one of them extending into the Bahamas.
Lindroth (1968: 934-938, as
This species ranges from Massachusetts (Suffolk County, MCZ) to central Iowa (Wickham 1911b: 8), including southernmost Ontario (Lindroth 1968: 935; Bousquet 1987a: 132), south to southern Texas (Johnson 1978: 67) and central Florida (Peck and Thomas 1998: 21); also recorded from north-central Maine (Majka et al. 2011: 46). The records from Colorado (Elias 1987: 634) and southeastern Arizona (Dajoz 2004: 116) need confirmation.
This species is found from Maine (Procter 1946: 111) to South Dakota (Kirk and Balsbaugh 1975: 33), including southern Quebec (Larochelle 1975: 31) and southern Ontario (Lindroth 1968: 937), south to southeastern Colorado (Miller and Peairs 2008: 34), southeastern Texas (Casey 1914: 288), and northeastern Florida (Saint Johns County, MCZ).
This species occurs from Nova Scotia (Lindroth 1954c: 309) to eastern South Dakota (Kirk and Balsbaugh 1975: 33; French et al. 2004: 557), south to southeastern Colorado (Miller and Peairs 2008: 34), southern Texas (Wickham 1897: 113; Harris County, Foster F. Purrington pers. comm. 2009), and southern Florida (Monroe County, Drew A. Hildebrandt pers. comm. 2007).
This species ranges from southern Quebec (Larochelle 1975: 32) to southeastern South Dakota (Kirk and Balsbaugh 1975: 33), south to eastern Texas (Snow 1906a: 141, as
Thirty-two species in the Nearctic (two western species), Australian (five species), Oriental (five species), Palaearctic (15 species), and Afrotropical (eight species) Regions.
The two North American species have never been compared and the specific independence of the two not tested.
This taxon is considered a distinct genus by several authors, including Jaeger and Kataev (2003: 399). However, the main character state separating members of
This species is known from a few localities from northwestern Oregon (Hatch 1953: 185) to southeastern Oregon (Harney County, CNC).
This species is known from California, from Humboldt County (CAS) to Santa Clara County (LeConte, 1857c: 29).
Four North American species, one of them extending into the Bahamas, placed in two subgenera.
Lindroth (1968: 925-926) treated all currently recognized species in his key to Canadian
This taxon has been treated as a subgenus of
One species occurring over eastern North America.
This species ranges from southern Quebec (Larochelle 1975: 31) south to eastern Oklahoma (Latimer County, UASM), northwestern Arkansas (Newton County, Peter W. Messer pers. comm. 2008), southeastern Mississippi (George and Greene Counties, Drew A. Hildebrandt pers. comm. 2008), east-central Kentucky (Jessamine County, MCZ), and northern Virginia (Hoffman and Roble 2000: 38).
Three eastern North American species of which one extends into the Bahamas.
This subgenus is in need of a taxonomic revision. I have seen at least four species in eastern North America, one of them (from Baker and Ware Counties in Georgia and Highlands County in Florida) being undescribed.
This species ranges from Georgia (LeConte 1869a: 378; Fattig 1949: 53) to eastern Texas along the Gulf Coast (Casey 1914: 264, as
This species ranges from southern Virginia (Hoffman and Roble 2000: 38) and Tennessee (Bradley, Crockett, Loudon, Maury, and Smith Counties, CMNH) to southern Florida (Peck and Thomas 1998: 21) and the Bahamas (Darlington 1953: 11), west at least to east-central Texas (Riley 2011), including southeastern Louisiana (Saint Tammany and Tangipahoa Parishes, CNC). The records from Rhode Island (Davis 1904: 14), Delaware (Houghton 1905: 212), and “Arkansas” (Bousquet and Larochelle 1993: 228) need confirmation.
The range of this species extends from southern Quebec (Larochelle 1975: 32) to eastern South Dakota (Kirk and Balsbaugh 1975: 33), south to eastern Texas (San Augustine and Hardin Counties, CMNH, UASM) and southern Florida (Peck and Thomas 1998: 21).
Two species in southern United States and Middle America (one species) and the West Indies (one species in Haiti).
Darlington (1936c: 205) discussed the structural differences between the two species.
This species is known from central (Grady County, Robert L. Davidson pers. comm. 2008) and southwestern Oklahoma (Kondratieff et al. 2005: 173), Arkansas (Garland and Ouachita Counties, Robert L. Davidson pers. comm. 2012), western Louisiana (Sabine Parish, CMNH), and southern Texas (Wickham 1897: 113) south at least to Nicaragua (Blackwelder 1944: 48).
About 25 species in the Western Hemisphere arrayed in two subgenera:
Nègre (1963) revised the species, including all three found in North America, known at the time.
Two species in southern North America and Mexico.
This species ranges from southern Arizona southwards through the Sonoran and Chihuahuan deserts to Oaxaca in Mexico (Ball and Shpeley 1992a: 55). The records from southwestern New Mexico (Fall and Cockerell 1907: 161) and “California” (Csiki 1932a: 1061) need confirmation.
This species is known from southern Arizona (Nègre 1963: 215) to western Texas (Culberson, Jeff Davis, and Presidio Counties, CMNH), south to Durango and Sinaloa (Nègre 1963: 215).
Twenty-three species in the Nearctic (one species) and Neotropical (23 species) Regions.
This species is known from southern Arizona to the state of Guerrero in western Mexico (Nègre 1963: 218).
Worldwide, with about 1,665 species (Lorenz 2005: 362-387, as
Noonan (1976) arrayed the genera of this subtribe into the following genus-groups: Acinopi (no North American representatives), Amblystomi (no North American representatives), Bleusei (no North American representatives), Bradybaeni (no North American representatives), Dapti (including
One North American species in the temperate regions.
Lindroth (1968: 745-746) treated the species.
This species ranges from Saskatchewan to central British Columbia (Lindroth 1968: 745-746), south to southern Arizona (Snow 1907: 142; Pima County, CMNH), central New Mexico (Fall and Cockerell 1907: 161; Bernalillo and Socorro Counties, CMNH, UASM), and southeastern Kansas (Wilson County, MCZ). One old specimen labeled “Tex” is known (MCZ).
One North American species in the temperate regions.
Ball (1960b) and Lindroth (1968: 747-748) treated the species.
This species ranges from western New York (Ball 1960a: 59, Fig. 3) to the Okanagan Valley in British Columbia (Lindroth 1968: 747), south to northeastern Oregon (Hatch 1953: 165), southern Arizona, southern Texas (Ball 1960a: Fig. 3), southern Louisiana (Allen 1965: 69), and northwestern Mississippi (Bolivar County, Drew A. Hildebrandt pers. comm. 2008), east to southern Georgia (Torres and Ruberson 2006: 32). The records from southeastern Pennsylvania (Rathvon 1869: 526, as
About 70 Palaearctic species arrayed in six subgenera:
Lindroth (1968: 756-758, as
About 40 Palaearctic species, of which a single one,
This Palaearctic species is adventive in North America where it is known from Prince Edward Island and Nova Scotia (Majka et al. 2006: 606) to eastern Iowa (Linn County, Doug A. Veal pers. comm. 2009), south to southern Pennsylvania [see Larochelle and Larivière 1989b: Fig. 1] and New Jersey (Steffens and Davidson 1979: 64). The first inventoried specimen collected on this continent was found in Long Island, New York, in 1954 (Dietrich 1958: 46).
This Palaearctic species is adventive in North America where it is known from Montreal (Lindroth 1968: 758) and the Quebec City region (Landry and Rancourt 1976: 53; CNC) in Quebec. The first inventoried specimen collected on this continent was found in Montreal in 1953.
About 415 species (Lorenz 2005: 363-372) in the Nearctic, Neotropical (Mexico only), Australian (one adventive species in New Zealand), Oriental, Palaearctic, and Afrotropical Regions. The Northern Hemisphere is represented by about 360 species (roughly 87% of the world fauna) and North America alone by 59 species (approximately 14%). Five species are Holarctic and three North American species are adventive.
About 60 species in North America (13 species), Mexico (four species, one of them,
Ball and Anderson (1962) revised the North American species and provided a key for their identification. Lindroth (1968: 758-765) covered ten species. Subsequently one new species was described by Will (2002a).
This species ranges from New Jersey and Virginia to northeastern North Dakota, south to the Gulf Coast of Texas and northwestern Mississippi (Bolivar County, Drew A. Hildebrandt pers. comm. 2007); also known from north-central Colorado [see Ball and Anderson 1962: Fig. 39].
This species ranges from Nova Scotia to southern Manitoba, south to southeastern Arizona, southeastern Texas, and central Florida [see Ball and Anderson 1962: Fig. 39].
This species is widely distributed east of the Rocky Mountains from Nova Scotia (Lindroth 1968: 764) to southern Manitoba, south to north-central Colorado, southern Oklahoma, and central Florida [see Ball and Anderson 1962: Fig. 39]. The record from south-central Montana (Hatch 1933a: 10) is probably in error.
This primarily prairie species ranges from southeastern Michigan to southern Alberta, south to central Arizona and the Rio Grande Basin in south-central Texas [see Ball and Anderson 1962: Fig. 39]. According to Ball and Anderson (1962: 76), the species is also found in “Mexico.” The records from “Arkansas” and “Pennsylvania” (Bousquet and Larochelle 1993: 232) need confirmation.
This eastern species occurs from the Saguenay River in southern Quebec to eastern South Dakota, south to central Kansas, southern Mississippi, and southwestern Georgia [see Ball and Anderson 1962: Fig. 39]. The record from “Nova Scotia” (Bousquet and Larochelle 1993: 231) was based on a misidentified specimen of
This species is known only from “Arizona” (Erwin et al. 1977: 4.48) and “Mexico” (Bates 1882a: 57). The record from “near Philadelphia,” Pennsylvania (Casey 1914: 82) is likely in error.
Ball and Anderson (1962: 45) listed
This common species occurs over a large area in North America from Prince Edward Island (Lindroth 1968: 761) and Nova Scotia to south-central British Columbia, south to southern California, northwestern Mexico, southeastern Texas, southern Florida, and the Bahamas [see Ball and Anderson 1962: Fig. 38].
This species is found from Maryland to southern Nebraska (Nuckolls County, Foster F. Purrington pers. comm. 2011), south to south-central Oklahoma, northern Louisiana, and east-central Alabama [see Ball and Anderson 1962: Fig. 38].
This species is restricted to the Coastal Plain ranging from south-central North Carolina to southern Florida, west to eastern Texas [see Ball and Anderson 1962: Fig. 38].
This species ranges from southwestern Maine (Majka et al. 2011: 46) and southern Quebec (Larochelle 1975: 87) to western North Dakota, south to western (Dajoz 2007: 23) and northeastern Texas, southeastern Louisiana, and the Florida Panhandle (Peck and Thomas 1998: 21); also known from northern (Villa-Castillo and Wagner 2002: 246) and east-central Arizona [see Ball and Anderson 1962: Fig. 37].
This species is known only from three specimens collected in Kansas (Ball and Anderson 1962: 43) and five specimens from central Oklahoma (Seminole County, Robert L. Davidson pers. comm. 2008).
This adventive species is found from Newfoundland (Larson and Langor 1982: 593) to near Montreal, Quebec (Mercado Cárdenas and Buddle 2007: 140), south to Connecticut (Krinsky and Oliver 2001: 210) and Rhode Island (Zhang et al. 1994: 69). The record from Wayne County, New York (Hajek et al. 2007) is in error (see Hajek et al. 2009: 913). The first inventoried specimen collected on this continent was found on Prince Edward Island in 1937 (Brown 1950b: 199).
This species is known only from two specimens collected in 1963 in two localities in Florida, along the coast of the Gulf of Mexico [see Will 2002a: Fig. 5].
Two North American species, both extending into northern Mexico.
Will (1997) reviewed the species of this subgenus and provided a key for their identification.
This taxon is listed as a synonym of
This species ranges from Nova Scotia (King County, CNC) to Washington, north to southern Manitoba (Lindroth 1968: 765) and southern Saskatchewan (CNC), south to the northern part of the Baja California Peninsula, northeastern Mexico, and northern Florida [see Will 1997: Fig. 1A].
This species ranges from northern Colorado (Will 1997: 47) to southern Virginia (Hoffman and Roble 2000: 37), north to Wisconsin (Purrington et al. 2002: 201), south to northern Florida, northeastern Mexico (Will 1997: 47, 49), southeastern New Mexico (Dunn 1986: 2), and southeastern Arizona (Cochise County, Ken Karns pers. comm. 2009) [see Will 1997: Fig. 1B].
Two North American species.
El-Moursy (1959) and Noonan (1991: 131-135) revised the species. Both species were also treated by Lindroth (1968: 766-767).
This species occurs east of the Rocky Mountains ranging from southern Quebec to southeastern Alberta, south to central New Mexico, southern Kansas, central Alabama [see Noonan 1991: Fig. 285], and southeastern South Carolina (Ciegler 2000: 100).
This species is restricted to mountains in “Utah” (Noonan 1991: 135), Arizona, New Mexico, and southern Colorado [see Noonan 1991: Fig. 285]. The record from “Mexico” (Csiki 1932a: 1186) is probably in error.
This group is represented in the Nearctic and Palaearctic Regions but the number of species cannot be assessed at this time. The North American fauna has 19 species.
The species of this subgenus have been revised by Noonan (1991: 45-127). Lindroth (1968) covered all the species except
This taxon corresponds to the
This species is known only from the type locality in east-central New Mexico.
This species is restricted to arid grasslands in eastern Arizona, southern Colorado, and New Mexico [see Noonan 1991: Fig. 276].
Ball (1972: 185) proposed
This species ranges from the Alexander Archipelago and northwestern British Columbia south to southwestern Oregon, east to eastern Washington [see Noonan 1991: Fig. 276].
This western species occurs from south-central British Columbia to western Minnesota (Gandhi et al. 2005: 930), south to east-central and westernmost Texas, southern Arizona, and northern California [see Noonan 1991: Figs 279-282].
Noonan (1991: 119-124) recognized two morphs within this species, the
This species is found along the Coastal Plain and Piedmont Plateau from southeastern New York and New Jersey to northern Florida, west to southeastern Oklahoma and southeastern Texas [see Noonan 1991: Fig. 270]. The record from southwestern Ohio (Blatchley 1910: 186) is probably in error.
This species ranges from southernmost Ontario (Bousquet 1987a: 130) to southeastern South Dakota (Kirk and Balsbaugh 1975: 28), south to southeastern Kansas (Noonan 1991: 115) and northern Mississippi (Drew A. Hildebrandt pers. comm. 2007). Two old specimens simply labeled from Louisiana and Florida are known (Noonan 1991: 115).
This western species occurs from northwestern British Columbia to southwestern Saskatchewan, south to north-central New Mexico and northern Arizona in the Rocky Mountains and to northern Oregon along the coast [see Noonan 1991: Fig. 267]. The record from south-central South Dakota (Kirk and Balsbaugh 1975: 29, as
This species is transcontinental in the north ranging from Newfoundland (Lindroth 1955a: 139; probably only as strays) to western Yukon Territory, south to northern Oregon along the west coast, southeastern Arizona and central New Mexico in the Rocky Mountains, the Black Hills in western South Dakota, northern Illinois, and southwestern Pennsylvania (Noonan 1991: 56) in the east [see Noonan 1991: Fig. 268]. The records from Connecticut (Britton 1920: 218; see Krinsky and Oliver 2001: 5), Indiana (Blatchley 1910: 186), and southwestern Oklahoma (Kondratieff et al. 2005: 172) need confirmation.
This eastern species ranges from southern Quebec to northern Minnesota, south to central Missouri and southwestern Virginia (Hoffman and Roble 2000: 38) [see Noonan 1991: Fig. 267]; also known from southwestern South Dakota (Larsen and Purrington 2010: 571). An old specimen simply labeled from Kansas and one labeled from Alabama are known (Noonan 1991: 51).
This species ranges from Maine (Piscataquis County, CNC) and northern New Brunswick (Webster and Bousquet 2008: 20) west to Manitoba and Minnesota (Lindroth 1968: 785), south to the Black Hills in western South Dakota (CMNH). The record from the Ungava Bay in Labrador (Sherman 1910: 181) is probably in error.
This species is externally very similar to
This Holarctic species ranges in the Nearctic Region from Newfoundland to the Alaskan coast, south to the Sierra Nevada in California, southern Arizona, southeastern Nebraska, and New Hampshire and New York [see Noonan 1991: Figs 277 and 278]. The species is known only from a few scattered localities in the prairies. The record from southern Indiana (Wiedenmann et al. 1992: 286) needs confirmation. In the Palaearctic Region, the species is known from Sweden to the Far East and the Altai in Kazakhstan (Kataev et al. 2003: 379).
Noonan (1991: 102-104) recognized two morphs in this species, the
This western species occurs from Vancouver Island to southeastern Manitoba (Roughley et al. 2010: 230), north to northeastern British Columbia, south to western Texas, southern Arizona, and the White Mountains in California, east to eastern Nebraska [see Noonan 1991: Fig. 271].
Noonan (1991: 75) recognized two forms within this species, a southern one, known as
This species is transcontinental in the north, ranging from Newfoundland to the Aleutian Islands in Alaska, south to the Lillooet Land District in British Columbia, north-central New Mexico along the Rocky Mountains, the Black Hills in western South Dakota, and New England in the east [see Noonan 1991: Fig. 276].
This species ranges from the Nova Scotia Peninsula (Christopher G. Majka pers. comm. 2008) to southern South Dakota (Kirk and Balsbaugh 1975: 30), north to southeastern Manitoba (Roughley et al. 2010: 230), south to Nebraska and southern Pennsylvania (Noonan 1991: 85; Fig. 272).
This Holarctic species ranges in North America from Newfoundland to southeastern Alaska (Lindroth 1968: 777), south to west-central Oregon, southern Nevada, southeastern Arizona and south-central New Mexico along the Rocky Mountains, the lower peninsula of Michigan, and southern New York [see Noonan 1991: Fig. 269]. The records from southern Indiana (Wiedenmann et al. 1992: 282, as
This species ranges from Nova Scotia to central Alberta and southeastern Northwest Territories, south to southern Wisconsin and Connecticut [see Noonan 1991: Fig. 270]. All known sites in the United States are east of the Mississippi.
With the exception of two specimens labeled from “British Columbia” and “Victoria” on Vancouver Island (Noonan 1991: 70), this species occurs east of the Rocky Mountains from central Alberta to the Quebec City area (Larochelle 1975: 87, map 302), south to Massachusetts, northwestern Pennsylvania (Erie County, Robert L. Davidson pers. comm. 2012), southern Kansas, central New Mexico, and southeastern Arizona [see Noonan 1991: Fig. 271]. This species is much more abundant in the prairies of central North America and both Lindroth (1968: 779; 1971: 1456, 1457) and Noonan (1991: 69) concluded that the species was originally an inhabitant of the prairies that expanded its range eastwards after humans cut down forests in the east.
This eastern species ranges from southern Quebec, southern Ontario, and the New England states south through the Appalachian mountains to northern Georgia [see Noonan 1991: Figs 273-275]. The record from southwestern Manitoba (Stjernberg 2011: 70) is probably in error.
Noonan (1991: 87-88) recognized two morphs for this species, the
This species ranges from central Alberta to southeastern Manitoba (Roughley et al. 2010: 230), south to western Kansas, northern New Mexico, and northeastern Utah [see Noonan 1991: Fig. 272].
This subgenus includes native species in North America, Mexico, and the Palaearctic and Afrotropical Regions. The total number of species cannot be determined at this time. The North American fauna is represented by 17 species, of which two are adventive.
Noonan (1991) revised all the North American species except the two adventive ones but his key (pages 20-45) included all 17 species. Lindroth (1968) covered all but three (
This Palaearctic species is adventive in North America where it is known in the east from Newfoundland (Lindroth 1955a: 137) to eastern Minnesota (Ramsey County, CMNH), south to “Kansas” (Lindroth 1955a: 137) and northern (Tucker and Preston Counties, CMNH) and eastern West Virginia (Pendleton County, Foster F. Purrington pers. comm. 2009). The record from southern Florida (Peck and Thomas 1998: 21) may represent a separate introduction. In the west, the species ranges from Alberta and British Columbia, including Vancouver Island (Lindroth 1968: 769), south to western Oregon (Clackamas, Linn, and Yamhill Counties, CMNH) and southwestern Idaho (Owyhee County, Ken Karns pers. comm. 2009); the species is also found in southeastern Arizona (Cochise County, CNC). The first inventoried specimen caught in the east was collected prior to 1798 in Pennsylvania (Palisot de Beauvois, 1811: 108, as
This Palaearctic species is adventive in North America where it is known from Nova Scotia (Majka et al. 2006: 606) and Prince Edward Island (Majka et al. 2008: 132) to eastern Ontario (near Reids Mills, CNC), including southern Quebec (Chantal 1994: 29), south to eastern Pennsylvania (Davidson et al. 2011: 517). The first inventoried specimen collected on this continent was found in New Hampshire in 1981 (Bell and Davidson 1987: 56).
This species is found west of the Mississippi River from northeastern Alaska to northwestern Minnesota, south to the highlands of central Mexico (Noonan 1991: 179) and southern California [see Noonan 1991: Fig. 289].
Three subspecies of
This species is known only from a few scattered localities from southwestern British Columbia, including Vancouver Island, south to west-central Oregon, east to northern Idaho [see Noonan 1991: Fig. 291].
This species is known from only six specimens collected near the coast in Massachusetts and New Jersey [see Noonan 1991: Fig. 283]. The collection date of the last known specimen is 1932 and Noonan (1991: 153) believes that it “might be extinct because of destruction of ocean side habitats.”
This western species occurs from central Alaska south to southern California, southeastern Arizona, and south-central Colorado, east to western Nebraska [see Noonan 1991: Fig. 286]. The records from northeastern Kansas (Popenoe 1877: 24) and Beaver Island in Michigan (Hatch 1925: 554) are probably in error. According to Noonan (1991: 143), this species is one of the most abundant
The range of this species extends from central Alberta and southern Saskatchewan south to southwestern Texas and southeastern Arizona; seemingly isolated in Minnesota (Gandhi et al. 2005: 930) and northwestern Oregon [see Noonan 1991: Fig. 287]; also recorded from northern Sonora in Mexico (Bates 1884: 271).
This species ranges from western Quebec to Vancouver Island, north to southern Northwest Territories, south to the Sierra Nevada in California, southern Colorado in the Rocky Mountains, southwestern South Dakota (Custer County, Peter W. Messer pers. comm. 2011), the upper peninsula of Michigan, and southwestern Pennsylvania [see Noonan 1991: Fig. 288]. Several specimens simply labeled from “New Mexico” are known (Noonan 1991: 159). East of the Rocky Mountains, the species is known only from a few scattered localities.
The range of this species extends from Alaska (Lindroth 1968: 808) east to Anticosti Island in Quebec, south to northeastern Minnesota (Purrington and Maxey 2007: 219) and the southern part of the Prairie Provinces [see Noonan 1991: Fig. 287]. The records from Colorado (Wickham 1902: 242) and New Mexico (Fall and Cockerell 1907: 161) are likely in error.
This Holarctic subspecies ranges from eastern Siberia (Kataev et al. 2003: 384) to the Alaska-Yukon border along the Porcupine River. On this continent, the subspecies is known from extant specimens only from the type locality. It was cited as a dominant taxon in a late Pleistocene site in the Klondike region of Yukon Territory (Zazula et al. 2006: 265).
This taxon was originally described as a distinct species but Kataev (1990: 396) listed it as a subspecies of
This species ranges from southwestern Alberta and British Columbia (Noonan 1991: 162-163) south to west-central Oregon, northeastern Nevada, and northwestern New Mexico (Casey 1924: 106, as
This species is known from scattered localities from Newfoundland (Lindroth 1955a: 143) to southern British Columbia, north to southwestern Northwest Territories, south to central Arizona, north-central New Mexico, eastern Texas, and southwestern North Carolina [see Noonan 1991: Fig. 292].
This species ranges from southern New Brunswick (Webster and DeMerchant 2012: 6) to south-central Alaska (Lindroth 1968: 806), south to the Sierra Nevada in California, central Arizona and central New Mexico along the Rocky Mountains, southern Indiana, and Maryland (Erwin 1981b: 175) [see Noonan 1991: Fig. 296].
The range of this species extends from Nova Scotia to south-central British Columbia, north to central Alberta, south to southern Oregon, east-central Arizona and central New Mexico along the Rocky Mountains, the Texas Panhandle (Michels et al. 2010: 743), northern Missouri, and southern Georgia [see Noonan 1991: Fig. 293]. The record from “Florida” (Leng 1920: 71) needs confirmation.
This species is known from only 16 specimens (including 15 ♂) from mountains in southern California [see Noonan 1991: Fig. 287] and could be either endangered or extinct according to Noonan (1991: 189). The last known specimen found was collected in 1919.
The range of this Holarctic species extends in northern North America from Newfoundland to south-central Alaska (Lindroth 1968: 799), south to east-central British Columbia, “Minnesota” (Lindroth 1968: 799), northeastern Wisconsin (Messer 2010: 41), “Michigan” (Lindroth 1968: 799), and Maine (Noonan 1991: 187) [see Noonan 1991: Fig. 294]. The records from the Sierra Nevada and Colorado (Elias 1987: 633) are probably in error; that from Prince Edward Island (Bousquet and Larochelle 1993: 237) needs confirmation (see Majka et al. 2008: 132).
This species is transcontinental in the north ranging from Newfoundland to the Kenai Peninsula in Alaska, south to southern California, central Arizona and New Mexico in the Rocky Mountains, Nebraska, and northern Virginia [see Noonan 1991: Figs 297-309]. The records from central Alabama (see Noonan 1991: Figs 302 and 304), “Arkansas” (Bousquet and Larochelle 1993: 237) and southern Kansas (Snow 1903: 194, as
Noonan (1991: 196-207) recognized six morphs within this species, the
Six species in southwestern North America.
Ball (1972) revised the species and provided a key for the identification of the males.
This species is known only from western Texas (Ball 1972: Fig. 13) and southeastern New Mexico (Perrault 1974: 120).
This species is known for sure only from Summit and Washington Counties in Utah (Ball 1972: 192). The record from “Arizona” (Bousquet and Larochelle 1993: 237) needs confirmation.
This species is known only from the holotype collected in southeastern Arizona [see Ball 1972: Fig. 13].
This species is found in southern (Perrault 1974: 120) and western New Mexico and southeastern Arizona [see Ball 1972: Fig. 13].
This species is known only from northwestern Arizona [see Ball 1972: Fig. 13; Perrault 1982b: 269].
Ball (1973: 74) proposed
This species is known only from Culberson County in western Texas and Pima County in southern Arizona [Ball 1972: 194, Fig. 13].
Four Palaearctic species, of which one is Holarctic.
The North American species was covered in Lindroth (1968: 815-817) monograph under the name
This Holarctic species ranges from Germany to eastern Siberia (Kataev et al. 2003: 376) and from Alaska to Nova Scotia (Lindroth 1968: 817, as
One Holarctic species.
The species was treated by Lindroth (1968: 748-749) and Noonan (1991: 212-213).
Noonan (1991: 211) treated this taxon as a group within the genus
This Holarctic species ranges from the European part of Russia to eastern Siberia (Kataev et al. 2003: 370) and in the Nearctic Region from east-central Alaska to southern Northwest Territories (Lindroth 1968: 749), south to central Alberta (Bousquet 1987a: 130). The species is rarely collected on this continent.
Two allopatric species in temperate areas of North America.
Ball (1976b) revised the species and provided a key for their recognition.
This species is known from “Arkansas” to eastern New Mexico [see Ball 1976b: Fig. 7], including southeastern Colorado (Michels et al. 2008).
This species in found along the Mississippi Basin from west-central Wisconsin (Messer 2010: 41) to eastern Nebraska (Colfax County, Foster F. Purrington pers. comm. 2010; Greeley County, R. Michael Brattain collection), south to Oklahoma (Ball and Bousquet 2000: 99) and Missouri [see Ball 1976b: Fig. 7].
Twenty-four species in temperate, subtropical, and tropical areas of the Nearctic (four species, only one endemic) and Neotropical (23 species) Regions, including the West Indies (three species, one of them endemic).
Ball and Maddison (1987) revised the species and provided a key for their identification.
This species ranges from southeastern California and southwestern Arizona, south along the Pacific Coast to El Salvador [see Ball and Maddison 1987: Fig. 29].
This species is found along and near the Atlantic Coastal Plain from New Jersey to the Florida Keys [see Ball and Maddison 1987: Fig. 28] and along the Gulf of Mexico in southwestern Alabama (Löding 1945: 25; Robert L. Davidson pers. comm. 2012). The record from “Pennsylvania” (LeConte 1847: 392), “Illinois,” and “Texas” (Horn 1880e: 182) are probably in error.
This species ranges along the Coastal Plain from New Jersey to southern Florida, west to southeastern Texas, north along the Mississippi Basin to southern Arkansas (Ouachita County, CMNH) and northeastern Mississippi; south of the Rio Grande it occurs through eastern Mexico and Central America to the Canal Zone in Panama [see Ball and Maddison 1987: Fig. 29]. The record from “Pennsylvania” (Ball and Maddison 1987: 210) needs confirmation.
The range of this species extends along the Gulf Coast from the Florida Panhandle (Jefferson County, CMNH) to central Veracruz in Mexico [see Ball and Maddison 1987: Fig. 31].
Eighteen species in the Neotropical Region, of which one extends into southern Texas.
There is no modern key for the identification of the species. The genus is in need of a revision.
This species is known from northeastern Arkansas (Kraim 1983: 137, as
The systematic position of this species is not quite settled. It was transferred to the genus
Western Hemisphere, with about 190 species arrayed in two subgenera:
Putzeys (1878) reviewed the genus but his work is outdated. About one-third of the currently recognized valid North American species were described by Casey and there is little doubt that several of them will turn out to be synonyms. A revision of the genus is much needed.
Fourteen species in temperate, subtropical, and tropical areas of the Nearctic (nine species) and Neotropical (eight species) Regions.
This species is known only from the type locality in southeastern Texas.
This species is known from North Carolina (Casey 1914: 144) and northeastern Georgia (Fattig 1949: 48).
This species is found in Mississippi (Harrison, Jackson, and Stone Counties, Paul K. Lago pers. comm. 2009), throughout the Florida Peninsula (Peck and Thomas 1998: 22), on several islands of the West Indies (Peck 2005: 32; Peck 2011: 13), and in Venezuela (Reiche 1843b: 142, as
The range of this species extends from west-central Maine (Majka et al. 2011: 46) to eastern Minnesota (Gandhi et al. 2005: 931), south to “Texas” (Horn 1880e: 183) and southern Florida (Peck and Thomas 1998: 22); also cited from “northern Mexico” (Casey 1914: 143).
This species is known from North Carolina (Brimley 1938: 128), Georgia (Horn 1880e: 182; Fattig 1949: 48), throughout Florida (Peck and Thomas 1998: 22), Alabama (Löding 1945: 25), and southeastern coastal Mississippi (Drew A. Hildebrandt pers. comm. 2007).
This species ranges from “Massachusetts” (Lindroth 1968: 830) to southern Florida (Vince Golia pers. comm. 2007), west to southern Texas (Johnson 1978: 67), and north along the Mississippi Basin to eastern Iowa (Linn County, Doug A. Veal pers. comm. 2009), “Illinois” (Bousquet and Larochelle 1993: 239), and “Indiana” (Schrock 1985: 355).
This species is known only from the type locality in central North Carolina.
This species is known only from the type locality in central North Carolina.
This species is known only from the type series.
About 175 species in temperate, subtropical, and tropical areas of the Nearctic (29 species) and Neotropical (about 150 species) Regions.
This species is known from northern Colorado (Denver and Boulder Counties, UASM) and southern Utah (Kane and Garfield Counties, CMNH) south to the Isthmus of Tehuantepec in Mexico (UASM).
This species is known from North Carolina (Manee 1915: 175), northeastern Georgia (Fattig 1949: 49), and southern South Carolina (Ciegler 2000: 103).
This species is known from a few localities on the lower Rio Grande of southern Texas (Purrington 2000: 9).
This species is known yet only from the Huachuca Mountains in southeastern Arizona.
This species is known only from the type series.
This species is known only from southeastern Texas.
This species ranges from “Georgia” (LeConte 1847: 408) to central Florida (Peck and Thomas 1998: 22), west to southwestern Alabama (Löding 1945: 25).
This species is known from southern Arizona (Cochise, Graham, Maricopa, Pima and Santa Cruz Counties, UASM), northern Sonora (UASM), Baja California Norte (UASM), and “California” (CMNH).
This species ranges from North Carolina (Brimley 1938: 128) and northern Tennessee (Montgomery County, Foster F. Purrington pers. comm. 2011) to southern Florida (Peck and Thomas 1998: 22), west to eastern Texas (Johnson 1978: 68) and south to eastern Mexico (George E. Ball pers. comm. 2008).
The range of this species extends from Nova Scotia (Kings County, NSNH) to central Wisconsin (Purrington et al. 2002: 201), south to central Texas (Travis County, UASM) and southern Florida (Peck and Thomas 1998: 22). The records from southern Arizona (Horn 1880e: 181; Wickham 1898: 301) need confirmation.
This species is known only from the type locality in central Texas.
This species is found from southwestern Maine (Majka et al. 2011: 46) to eastern Minnesota (Gandhi et al. 2005: 931), including southernmost Ontario (possibly only as strays) (Lindroth 1968: 821), south to eastern Texas (Nacogdoches County, CMNH) and central Florida (Manatee County, CNC).
This species is known from central Louisiana (Casey 1914: 151) and Texas (Leng 1920: 72; McCulloch County, USNM). One specimen determined by Casey (USNM) from Kansas is known.
This species is known from southern Florida and several islands of the West Indies (Peck and Thomas 1998: 22).
This species ranges from central Texas south at least to southern Tamaulipas in Mexico (LeConte 1858b: 59). The record from “Arizona” (Bousquet and Larochelle 1993: 241) needs confirmation.
Horn (1880e: 180) regarded
This species is known from central and southern Florida (Vince Golia pers. comm. 2007; Highland and Palm Beach Counties, CMNH, UASM), southern Mississippi (Hancock and Jackson Counties, Drew A. Hildebrandt pers. comm. 2008; Lago et al. 2002: 202), and southern Texas (Casey 1914: 148; Zapata and San Patricio Counties, CMNH, UASM).
The range of this species extends from western New Brunswick (Webster and Bousquet 2008: 19) to Minnesota (Epstein and Kulman 1990: 215) and South Dakota (Kirk and Balsbaugh 1975: 30), including southern Quebec and Ontario (Lindroth 1968: 824), north to southeastern Manitoba (Roughley et al. 2010: 230), south to southern Texas (Johnson 1978: 67), the Florida Keys (Peck and Thomas 1998: 22), and South
This species is known only from the type series.
This species is found throughout much of southern United States from North Carolina (Brimley 1938: 128) to the Florida Keys and the Bahamas (Peck and Thomas 1998: 22), west to southeastern California (Andrews et al. 1979: 28) and the Baja California Peninsula (Horn 1894: 312), including southern Illinois (Union and Pope Counties, UASM), south to southeastern Texas (Wickham 1897: 113; Johnson 1978: 68; Cameron and San Patricio Counties, UASM) and San Luis Potosí (UASM). The record from “Virginia” (Bousquet and Larochelle 1993: 241) needs confirmation.
This species is known from southern Florida (Peck and Thomas 1998: 22, as
This species is found from Connecticut (Krinsky and Oliver 2001: 218) to southern Montana (Hatch 1933a: 10), south to southern New Mexico (Fall and Cockerell 1907: 162), southern Texas (Johnson 1978: 67), and central Florida (Peck and Thomas 1998: 22), west along the south to southern Arizona (Griffith 1900: 566; Snow 1906b: 163; Snow 1907: 142; Greenlee and Pima Counties, CMNH) and Baja California (Horn 1894: 311).
This species ranges from southernmost Ontario (possibly only as strays) to south-central British Columbia (Lindroth 1968: 828), south to northeastern Oregon (Westcott et al. 2006: 9), Durango in Mexico (Ball and Shpeley 1992a: 59), northwestern Mississippi (Bolivar County, Drew A. Hildebrandt pers. comm. 2008), and northwestern Ohio (Lucas County, Harry J. Lee, Jr. pers. comm. 2008).
This species is known only from the type locality in western Mississippi.
This species is known from the Huachuca Mountains and Yuma County in southern Arizona (Schaeffer 1910: 403) and from Riverside County in southeastern California (Andrews et al. 1979: 28, as
This species is known only from the type series.
This species is known from the type locality in southern Arizona, from southern New Mexico (Dona Ana County, CMNH), and from southern Sonora in Mexico (UASM).
This species is known so far from Florida and several islands of the West Indies (Peck and Thomas 1998: 22; Peck 2009a: 13), from south-central Louisiana (Saint Landry Parish, Igor M. Sokolov pers. comm. 2009), as well as from southern Texas (Johnson 1978: 67; San Patricio and Cameron Counties, UASM), southern Mexico (Putzeys 1878: 33), and Colombia (Martínez 2003: 9). The record from Georgia (Fattig 1949: 48, as
This species is known only from the type locality in southern Florida.
This peculiar species is known from southern Kansas (Barber County, Robert L. Davidson pers. comm. 2012), west-central Oklahoma (Horn 1880e: 181), and northeastern Texas (Hunt County, CNC).
The systematic position of this species is not settled. It was not listed by Noonan (1985) in his classification and names of the Selenophori group. The species is markedly distinctive in its structural characters.
Twenty-eight species in North America (19 species) and Middle America (11 species).
This genus has never been revised and such study is much needed. More than 60% of the currently valid North American species have been described by Casey and some of them will certainly fall in synonymy.
This species is known from western Missouri (Vernon County, UASM), Texas (several Counties, UASM), southeastern New Mexico (Chaves County, UASM), and southeastern Arizona (Cochise County, UASM).
This species ranges from eastern Wyoming (Lavigne 1977: 45) to southeastern Oregon (Westcott et al. 2006: 8), south to southern California (Fall 1901a: 50) and south-central New Mexico (Fall and Cockerell 1907: 161).
This species is known from southeastern Colorado (Bent County, CNC), central New Mexico (Bernalillo County, CNC), and southern Arizona (Cochise and Santa Cruz Counties, UASM).
This species is known from southern Arizona (Horn 1891: 34; Griffith 1900: 565; Cochise and Maricopa Counties, UASM), New Mexico (Hidalgo and Quay Counties, CMNH; Luna County, UASM; Fall and Cockerell 1907: 161), western Texas (Brewster County, UASM), and Coahuila, Sonora, and Baja California Norte in Mexico.
This species is known, besides the type series, from a few specimens collected in Cochise County in southeastern Arizona (Robert L. Davidson, pers. comm. 2012).
This species is known from northern Texas only (Casey 1924: 120).
This species ranges from Kansas (Knaus 1907: 233; Trego, Scott, and Barber Counties, CMNH, CNC) to southeastern Utah (San Juan County, UASM), south to northern Sonora (Bates 1884: 276), Coahuila (UASM), and southeastern Texas (San Patricio County, UASM).
This species is known only from the type locality in central Texas.
This species is known only from Cochise County in southeastern Arizona (UASM).
This species is known from southeastern Arizona (Cochise County, UASM) and southern New Mexico (Casey 1924: 119).
This species ranges from southeastern New Hampshire (Rockingham County, Ross T. Bell pers. comm. 1992) to the Okanagan Valley in south-central British Columbia (Lindroth 1968: 831), south to northern Oregon (Union, Wallowa, and Wasco Counties, James R. LaBonte pers. comm. 2009), northern Utah (Salt Lake County, Foster F. Purrington pers. comm. 2009), New Mexico (Fall and Cockerell 1907: 161), western and central Texas (Lee and Brewster Counties, MCZ; Lindroth 1968: 831), southwestern Alabama (Clarke County, CMNH), central Georgia (Fattig 1949: 49), and eastern South Carolina (Ciegler 2000: 104).
This species is known only from the type locality in western Texas.
This species is known only from the type locality in western Texas.
This species is known from southern Arizona (Cochise County, UASM).
This subspecies is known only from the type locality in southeastern Arizona.
This subspecies ranges from southern Arizona (Wickham 1898: 301; Griffith 1900: 565; Snow 1906b: 162; Snow 1907: 142) and northern Sonora in Mexico (Bates 1884: 276, as
This species is known from Arizona (Cochise, Gila, Graham, Pima, Santa Cruz, and Yavapai Counties, UASM; Casey 1914: 160) and western Texas (Terrell and Jeff Davis Counties, UASM).
This species is known only from the type series.
This species has been recorded only from Santa Fe in New Mexico (Fall and Cockerell 1907: 161).
This species is known only from the type locality in northeastern Texas.
Six species in North America (three species), Middle and South America (five species), and the West Indies (two species, one of them endemic).
Ball et al. (1991) revised the species.
Ball et al. (1991: 942) postulated that the Neotropical genus
This subspecies is known from a small area in southeastern Arizona and southwestern New Mexico [see Ball et al. 1991: Fig. 24].
Besides the two subspecies found in North America, two other subspecies are known, one (
This subspecies ranges from northern Missouri to southern Nebraska (Adams County, Foster F. Purrington pers. comm. 2009), south to the Yucatán Peninsula, west to the southern portion of the Baja California Peninsula, east to central Alabama [see Ball et al. 1991: Fig. 24]. One specimen is known from Clemson, South Carolina (Ciegler 2000: 104).
This species ranges from southern Arizona southwards along the Pacific Versant of Mexico to central Jalisco [see Ball et al. 1991: Fig. 21].
This species ranges from southeastern Arizona and southwestern New Mexico south along the Pacific Versant of Mexico to Guerrero; it is also known from the southern parts of the Baja California Peninsula [see Ball et al. 1991: Fig. 23]. As pointed out by Ball et al. (1991: 951) the record from central Colorado based on a male is questionable but not impossible.
About 250 species (Lorenz 2005: 380-382), more than 85% of them inhabiting Asia, arrayed in six subgenera:
Lindroth (1968) reviewed all four North American species, one of them (
About 120 species in the Nearctic (two eastern species), Australian (six species), Oriental (about 25 species), and Palaearctic (about 90 species, the vast majority in its eastern part) Regions.
The range of this species extends from southern Quebec (Serge Laplante pers. comm. 2002) to southeastern South Dakota (Kirk and Balsbaugh 1975: 30), south to “Texas” (Lindroth 1968: 819) and central South Carolina (Ciegler 2000: 105).
Say (1830c: 19) noted that “
The range of this species extends from western New Brunswick (Webster and Bousquet 2008: 19) to southwestern Wisconsin (Grant County, Peter W. Messer pers. comm. 2008), south to northern Arkansas (Boone County, UASM) and eastern Georgia (Fattig 1949: 47). The records from “Florida” and “Minnesota” (Bousquet and Larochelle 1993: 244) need confirmation.
Six species in the Nearctic (two species) and east Palaearctic (four species) Regions.
Kataev (in Ball and Bousquet 2000: 96) revalidated
This species ranges from west-central Maine (Majka et al. 2011: 46) and southern Quebec (CNC) to east-central Minnesota (Gandhi et al. 2011: 673), north to Kapuskasing in central Ontario (CNC), south at least to northeastern Kansas (Popenoe 1877: 24), central Arkansas (Pulaski County, Robert L. Davidson pers. comm. 2008), southwestern Mississippi (Hinds County, CMNH), northern Georgia (Fattig 1949: 52), and eastern South Carolina (Ciegler 2000: 105).
This eastern species extends from Massachusetts (Middlesex County, Peter W. Messer pers. comm. 2008) to northeastern Kansas, including southernmost Ontario (Bousquet 1987a: 130) and southeastern Iowa, south to southeastern Texas and southern Florida, west to western Texas [see Noonan 1991: Fig. 286]. The record from southern Wisconsin (Rauterberg 1885: 21, as
This species was included in the genus
Nine species in eastern Mexico, one of them extends into southeastern Texas.
Ball (1970, 1976a) revised the species and provided a key (Ball 1976a: 62-63) for their identification.
This species is known only from southeastern Texas and northeastern Mexico (Ball 1970: 120).
One North American species in the temperate regions.
The species is covered in Lindroth’s (1968: 744-745) monograph.
The range of this species extends from Long Island, New York (Notman 1928: 244), to southeastern Alberta (Lindroth 1968: 745), south to southern Arizona (Cochise, Greenlee and Pima Counties, CNC, UASM), Durango (Bates 1891a: 241) and southern Coahuila (UASM) in Mexico, western Alabama (Pickens and Tuscaloosa Counties, UASM), and “Florida” (Leng 1920: 70, as
About 825 species in the Nearctic (13 species), Neotropical (16 species in Mexico), Oriental (about eight species), Palaearctic (about 775 species), and Afrotropical (17 species in Ethiopia) Regions, arrayed in six subtribes:
Several authors, including Lindroth (1966) and Ball and Bousquet (2000), placed the sphodrines within the tribe
About 100 species in Europe (about 60 species), northern Africa (about 20 species), western Asia, including India (about 25 species), and eastern North America (one species); most species occur in the Mediterranean region.
Lindroth (1968: 653) listed the genus
One species in the boreal and temperate regions of eastern North America.
Lindroth (1968: 653-654) treated the species in his monograph.
This species is found from Cape Breton Island (Bousquet 1987a: 128) to northern Minnesota (Gandhi et al. 2005: 929), south to southwestern North Carolina (Macon County, MCZ).
About 185 species (Lorenz 2005: 396-399) placed in the genus
About 185 species (Lorenz 2005: 396-399) in North America (eight species, one of them adventive), Mexico (14 species), Ethiopia (17 species), and the Palaearctic Region (about 145 species) arrayed in 11 subgenera:
Ball and Nègre (1972) revised the Western Hemisphere species and provided a key for their identification.
Fifty species (Lorenz 2005: 396-397) in Europe, the Middle East, and Nepal. Most of the species are endemic to the Mediterranean region. One species is adventive in western North America.
This Palaearctic species is adventive in North America where it is known from southwestern British Columbia (Lindroth 1966: 543) to northwestern Oregon (Westcott et al. 2006: 7). The first inventoried specimen collected on this continent was found in the vicinity of Vancouver, British Columbia, in 1928 (Hatch 1949c: 151).
Forty-five species in North America (six species), Mexico (13 species), and the Palaearctic Region (27 species, no endemic species in eastern Asia).
Gañán and Novoa (2006) considered
The record of
This species ranges from the southern Gulf Islands of southwestern British Columbia (James C. Bergdahl pers. comm. 1993) to southern Idaho, south to northern Utah and Mono County in the Sierra Nevada of California [see Ball and Nègre 1972: Fig. 53].
This species is found from Cape Breton Island in Nova Scotia (Lindroth 1966: 544) to western North Dakota (Tinerella 2003: 636), south to Nebraska, northern Alabama, and northeastern Georgia (Fattig 1949: 32) [see Ball and Nègre 1972: Fig. 51]. The records from Prince Edward Island (Bousquet and Larochelle 1993: 245, see Majka et al. 2008: 133) and eastern Kansas (Popenoe 1877: 23) need confirmation; those from “Florida,” Texas (Wickham 1896c: 133; Knaus 1905b: 348), and New Mexico (Fall and Cockerell 1907: 159) are likely in error (see Ball and Nègre 1972: 486).
The range of this transamerican species extends from Newfoundland (Lindroth 1955a: 112-113) to Alaska, including the Aleutian Islands (Lindroth 1966: 545), south to Washington (Hatch 1953: 131), central Arizona, southern New Mexico, the Black Hills in southwestern South Dakota (Kirk and Balsbaugh 1975: 23), and the Adirondack Mountains in northeastern New York (Notman 1928: 230) [see Ball and Nègre 1972: Fig. 53].
The range of this species extends from southern Quebec (Larochelle 1975: 69) to southeastern Utah, south to southeastern Arizona, southern Texas, and northern Florida [see Ball and Nègre 1972: Fig. 52]. The record from New Brunswick (see Majka et al. 2007: 11) is in error (Christopher G. Majka pers. comm. 2009).
This species is confined, as far as known, to mountains in southeastern Arizona (Ball and Nègre 1972: 505).
The range of this subspecies extends from southwestern British Columbia, including Vancouver Island, south to Humboldt County in northern California [see Ball and Nègre 1972: Fig. 40].
This subspecies occurs from eastern Washington and western Idaho south to central California [see Ball and Nègre 1972: Fig. 40].
This subspecies ranges from Mendocino County in California south to the northern parts of the Baja California Peninsula and southern Arizona (Ball and Nègre 1972: 482, Fig. 40); it also occurs on the Channel Islands. The record from “Nevada” (Ball and Nègre 1972: 482) is doubtful. The subspecies is adventive in Hawaii (Liebherr and Zimmerman 2000: 465).
According to Ball and Nègre (1972: 482), this form intergrades with the
Eleven species in North America (one species) and the Chinese provinces of Kansu, Szechwan, and Tsinghai, and the Autonomous Region of Tibet (ten species).
This taxon has been listed as a distinct genus in the subtribe
This species ranges in the east from eastern Newfoundland to Lake Superior, south to southwestern New York, and in the west from Kodiak Island in Alaska south to northern Oregon, east-central Nevada, and southern New Mexico along the Rocky Mountains, east to the Black Hills in southwestern South Dakota (Pennington County, CNC) [see Ball and Nègre 1972: Fig. 1]. The records from “California” (Lindroth 1955a: 114) and “Vermont” (Hamilton 1894a: 11) need confirmation.
About 100 species (Lorenz 2005: 400-401) in North America and Mexico (three species) and the Palaearctic Region (about 95 species), arrayed in four genera:
About 80 species in the Nearctic (two species), Neotropical (one species from northwestern Mexico), and Palaearctic (77 species, of which one is Eurasian, nine are from the Himalayas, and the remaining from eastern Asia) Regions.
Lindroth (1966: 550-552) treated both North American species.
This species is found in the southern part of the Rocky Mountains in western Utah (Casey 1924: 87, as
This species ranges from Newfoundland (Lindroth 1955a: 115-116) to southwestern British Columbia (Lindroth 1966: 551), south to western Washington (Hatch 1953: 132), northern Idaho (Hatten et al. 2011: 325), eastern Kansas (Cook and Holt 2006: 2313), and northern Georgia (Fattig 1949: 32). The record from southwestern Colorado (Wickham 1902: 238) needs confirmation.
About 360 species (Lorenz 2005: 401-407), all but one (
Casale (1988) revised all species then known.
About 195 species (Lorenz 2005: 403-406) in Europe, northern Africa, western and central Palaearctic Asia, and the Himalayas arrayed in 12 subgenera. Two species are adventive in several parts of the world.
Lindroth (1966: 549-550) treated both species found in North America.
Twenty species in Europe (eight species), northern Africa (four species), and Asia (13 species), one of them has been dispersed by trade to several continents and islands.
This species is now subcosmopolitan (Casale 1988: 460). It is adventive in North America and known from southern British Columbia, including Vancouver Island (Lindroth 1966: 550), to southwestern California, east across the Central Valley to the western foothills of the Sierra Nevada (David H. Kavanaugh pers. comm. 2008). The first inventoried specimen collected on this continent was found prior to 1874 since the species is reported by Crotch (1874: 12).
About 55 species in Europe, northern Africa, western Asia, and the Himalayas. One European species is adventive in North America and India (Casale 1988: 786-787).
This European subspecies is adventive in North America where it is known from Newfoundland (Lindroth 1966: 550; Larson and Langor 1982: 593) and the Maritimes (Lindroth 1966: 550) to the Saint Lawrence Estuary (Larochelle 1975: 98), from three specimens collected in 2005 and 2008 in Boston Harbor, Massachusetts (Davidson et al. 2011: 518), and from one specimen collected in 1980 in southeastern British Columbia (Bousquet 1987a: 125). The first inventoried specimen collected on this continent was found in Nova Scotia prior to 1894 (Bousquet 1992a: 507).
The subspecies
Worldwide, with about 2,670 species arrayed in 170 genera (Lorenz 2005: 407-437, excluding
Twenty-one species in temperate areas of the Nearctic (seven species) and Palaearctic (14 species, only one extending to eastern Asia) Regions.
Casey (1913: 169-171) published a key to all North American species except
This species has been recorded from Rhode Island (Casey 1913: 171) and from Richland County in South Carolina (Ciegler 2000: 106).
The range of this species extends from southern Quebec (Larochelle 1975: 96) to “Iowa” (Jaques and Redlinger 1946: 295), south to southern Louisiana (Calcasieu Parish, CNC) and northern Florida (Baker County, CMNH). The record from “Wisconsin” (Bousquet and Larochelle 1993: 248) needs confirmation.
This species is known only from the type series.
This species is known only from the holotype.
This species is known only from the original two specimens.
Lindroth (1966: 553) listed this form in synonymy with
This species ranges from “Nova Scotia” (Larochelle and Larivière 1990a: 30) to eastern Minnesota (Epstein and Kulman 1990: 215; Wickham 1896c: 134), south to northern Oklahoma (French et al. 2001: 229), Arkansas (Logan County, CNC), the Florida Panhandle (Leng 1915: 582), and southwestern Georgia (Fattig 1949: 35). The record from northern Colorado (Armin 1963: 158) needs confirmation.
This species is known only from the holotype collected in central Louisiana.
Eleven Mexican species, of which one extends into southern United States.
Liebherr (1991b) revised the species.
This species extends from east-central Arizona and southwestern Texas south to the state of San Luis Potosí in central Mexico [see Liebherr 1991b: Fig. 269]. Only two localities have been reported for the United States.
Eight species in North America (four species, only one of them endemic), Middle America (two species, no endemism), the West Indies (one endemic Cuban species), South America (one species, no endemism), Asia (four species, three of them endemic), and Europe (two species, no endemism).
Liebherr (1991b) revised the species. One species,
This species is widely distributed in the Western Hemisphere ranging from southeastern Manitoba to central Alaska, south to northern Oregon and through the Rocky Mountains, Transverse Volcanic, Central America, and the Andes to southern Ecuador, east in South America to western Venezuela [see Liebherr 1991b: Fig. 240].
This Holarctic species is found in the New World from west-central Alaska south to central California and to southern New Mexico along the Rocky Mountains. Disjunct populations are also known from Chiapas in Mexico and Guatemala [see Liebherr 1991b: Fig. 230]. In the Old World, the species is known from disjunct populations in Lapland, the Austrian Alps, and Burma [see Liebherr 1991b: Fig. 231].
This Nearctic species ranges from Newfoundland to Vancouver Island (Casey 1920: 97, as
This Holarctic species ranges in North America from central Alaska to Newfoundland (Lindroth 1955a: 117), south to eastern North Carolina and western Tennessee along the Appalachian Mountains, to northern New Mexico along the Rocky Mountains, and to central Washington [see Liebherr 1991b: Fig. 221]. In the Palaearctic Region, the range of the species extends from Great Britain to the Kamchatka Peninsula, south to the Philippines Islands, the Himalayan Mountains, and Italy in Europe [see Liebherr 1991b: Fig. 222].
One eastern North American species in the temperate regions.
Liebherr (1991b) redescribed the species and illustrated some of its structural character states.
This species is known from scattered localities in eastern North America ranging from western Vermont to “Minnesota,” south to east-central Texas (Riley 2011), southern Louisiana and central Florida [see Liebherr 1991b: Fig. 187]. The record from Sainte-Rose (= Laval) in Quebec (Hausen 1891b: 159) needs confirmation.
Fourteen species in the Nearctic (three species) and Palaearctic (ten species) Regions and one species (
Liebherr (1991b) revised the species except for three Palaearctic species of the subgenus
Eleven species in North America (three endemic species), Baja California (one endemic species), Asia (seven species, four of them endemic), Europe (three species), and northern Africa (one species).
This taxon is found from the Rocky Mountains in southwestern Alberta to south-central British Columbia, south to central Oregon and south-central Idaho [see Liebherr 1991b: Fig. 200].
This western species ranges from central Oregon to Baja California Norte, including west-central Nevada [see Liebherr 1991b: Fig. 200].
This western species is found from west-central British Columbia south to northern California [see Liebherr 1991b: Fig. 187].
Forty-nine species in North America (40 species) and Mexico (11 species) arrayed in six species groups.
Barr (1960b) revised the cavernicolous species and provided a key for their identification, but several new species have been described since. Barr (1974b) published a key to the species groups and a revision of the species of the
This species is known only from the type series collected in northwestern New Mexico.
This species is known from a few localities in southeastern Oregon (Harney County, James R. LaBonte pers. comm. 1992), southeastern Washington, and southern Idaho (Hatch 1953: 146; Stafford et al. 1986: 288). The record from northern Colorado (Armin 1963: 147) needs confirmation.
This species is known only from the type series collected in southeastern Arizona.
This species is known from “Nebraska” (LeConte 1863c: 8) to southwestern Colorado (LeConte 1879b: 54; Wickham 1902: 238), south to northern Arizona (Coconino County, CMNH; Snow 1906b: 162), the Sacramento Mountains in south-central New Mexico (Fall and Cockerell 1907: 159), and “Texas” (LeConte 1863c: 8).
This species is found from southern Saskatchewan and southern Alberta (Lindroth 1966: 647) south to Utah (Barr 1965b: 141), northeastern Colorado (Bell 1971: 52), and South Dakota (Kirk and Balsbaugh 1975: 25). The record from “Nebraska” (Bousquet and Larochelle 1993: 251) needs confirmation.
This species is known only from the type locality in northeastern New Mexico.
This species is known from many specimens collected in south-central Texas.
This species is known from a few specimens collected in southeastern Colorado and northwestern Texas (Barr 1960b: 50).
This species is known only from the syntype collected in central Colorado.
This species ranges from eastern Washington to southwestern Wyoming (Parmenter and MacMahon 1984: 26), south to northern Arizona (Wichkam 1896a: 157) and the Sierra Nevada in California (Casey 1913: 166). The records from Colorado (Cockerell 1893: 72; Wickham 1902: 238) need confirmation.
A preliminary study of the type specimens of
This species is known only from the type locality in southeastern Washington (Hatch 1953: 146).
This species ranges from southwestern Pennsylvania (Hamilton 1895: 350, 379; Ehrman 1900: 499) to southeastern Wisconsin (Rauterberg 1885: 14), south to northern Alabama (Jeannel 1949b: 95; Barr 1960b: 47) and northern Georgia (Fattig 1949: 32). The record from southwestern Colorado (Wickham 1902: 238) must be in error; that from “Arkansas” (Bousquet and Larochelle 1993: 250) needs confirmation.
This species is found from the Florida Panhandle (Choate and Rogers 1976: 364) to central Arkansas (Barr 1982a: 178), north to east-central Missouri (LeConte 1846b: 219). The records from northern Colorado (Wickham 1902: 238), northern Arizona (Wickham 1896a: 157), southern Montana (Hatch 1933a: 9), and southwestern Pennsylvania (Ehrman 1900: 500) are probably in error.
This species is known only from the original specimens collected in Fincher’s Cave, near Fayetteville, Arkansas (Barr 1960b: 49).
This species has been recorded from several localities in New Mexico (Fall and Cockerell 1907: 159; Casey 1913: 163).
This species is known from Greenlee (Casey 1913: 164) and Coconino Counties (CMNH) in southeastern and northern Arizona.
This species is known only from the type series collected in northwestern New Mexico.
This species is known only from the original specimens collected in the White Mountains, eastern California.
This species is known from a number of caves in Sutton, Edwards, and Pecos Counties, western Texas (Barr 1960b: 52).
This taxon has been considered a subspecies of
This species is known from a number of caves in Kerr, Edwards, Uvalde, and Bexar Counties, southern Texas (Barr and Lawrence 1960: 143; Reddell and Cokendolpher, 2004: 157).
This species is known only from the type locality in western Texas.
This species is known from numerous caves in Eddy County, southeastern New Mexico, and Culberson County, western Texas (Cokendolpher and Polyak 2004: 192).
This species has been recorded from southwestern Utah (Tanner 1928: 270), west-central (Bechtel et al. 1983: 474) and southern (Tanner and Tanner 1974: 218) Nevada, “Arizona” (Horn 1892c: 42), and New Mexico as far south as the Sierra Blanca Range (Fall and Cockerell 1907: 159).
This species is known from southern Arizona and western Chihuahua in northern Mexico (Barr 1982a: 182).
This species is found in several caves in central Travis County, Texas (Barr 1974b: 12).
This species is known from several caves on Camp Bullis, a U.S. Army installation in Bexar and Comal Counties, south-central Texas (Reddell and Cokendolpher, 2004: 154-155).
This species is known only from caves near Helotes and Camp Bullis in Bexar County, south-central Texas (Barr 1974b: 16; Reddell and Cokendolpher, 2004: 156).
This species has been listed as endangered by the U.S. Fish and Wildlife Service in December 2000.
This species is known from three caves in or near San Marcos, Hays County, in south-central Texas (Reddell and Dupérré 2009: 112).
This subspecies is known from three caves in southern Bexar County, south-central Texas (Reddell and Cokendolpher, 2004: 157).
This subspecies is known from a few small caves, near Helotes, in the highlands north of San Antonio, southern Texas (Barr 1974b: 23).
A series of eight specimens collected at Bat Cave in Government Canyon, Bexar County, Texas, are reported as hybrids
This species is known only from the type-locality cave in south-central Texas.
This species is known only from the type-locality cave in south-central Texas (Reddell and Cokendolpher, 2004: 161).
This subspecies is known from two caves in south-central Texas (Barr 1974b: 24).
This subspecies is known from a few caves in south-central Texas (Barr 1974b: 25).
This species is known only from Cobb Cavern and Cricket Cave in northern Williamson County, central Texas (Barr 1974b: 11).
This species has been collected in caves in northern Travis and southern Williamson Counties, central Texas (Reddell and Cokendolpher 2001: 110).
This species, also known under the vernacular name “Tooth Cave Ground Beetle”, is listed as an endangered species by the World Wildlife Fund.
This species is known from a number of caves on the Fort Hood Military Reservation in Bell and Coryell Counties, central Texas (Reddell and Cokendolpher 2001: 110).
This species is known only from the type-locality cave on the Jollyville plateau in central Texas.
This subspecies has been found yet only in two caves along Cibolo Creek in southern Texas (Barr 1974b: 16).
This subspecies is known only from two caves in south-central Texas (Barr 1974b: 15).
This subspecies is known from a few caves in Kendall and Comal Counties, southern Texas (Barr 1974b: 13; Reddell and Cokendolpher, 2004: 158).
This species is known yet only from the type-locality cave in south-central Texas (Reddell and Cokendolpher, 2004: 159).
This subspecies is known from several caves in central Texas (Barr 1974b: 8).
This subspecies is known from several caves in southern Williamson and northern Travis Counties, central Texas (Barr 1974b: 7).
This subspecies is found in a few caves in Real and Uvalde Counties, southwestern Texas (Barr 1974b: 20).
This subspecies is known from a few caves in Kerr, Real, and Bandera Counties, southwestern Texas (Barr 1974b: 19).
This species is known for sure only from the type locality in southwestern Nevada. The record from “California” (Bousquet and Larochelle 1993: 251) needs confirmation.
Eleven species in Texas (one species) and Mexico (ten species).
Barr (1982a) revised the species and provided a key for their identification.
Valentine (1987) believed this genus is closely related to the genera
This species is known from two nearby caves in southwestern Texas.
This species belongs to the
Five species in western North America (four species) and the Baja California Peninsula and Guadalupe Island in Mexico (two species, one of them endemic).
Liebherr (1985) revised the species. Subsequently he described a new species, the endemic Baja Californian
According to Liebherr (1989: 184), the genus
This species ranges through the Coast Ranges from central California south to the Mexican border [see Liebherr 1985: Fig. 27].
This species, also known under the vernacular name “tule Beetle”, ranges from southwestern Oregon south through California, including the Channel Islands, to Baja California Norte and Guadalupe Island [see Liebherr 1985: Fig. 25]. The species is apparently adventive and possibly established at Seattle, Washington (Liebherr 1985: 1194). The record from “Utah” (Bousquet and Larochelle 1993: 252) is likely in error.
This species occurs along the Coast Ranges of California from Siskiyou County to Santa Cruz and Alameda Counties and in the Sierra Nevada between Tulare and Shasta Counties [see Liebherr 1985: Fig. 27].
This species ranges from northwestern Oregon to northern California [see Liebherr 1985: Fig. 27]. An old specimen labeled from western Washington is known.
Two species endemic to the Canary Islands and one European species,
The North American species is covered in Lindroth’s (1966: 630-631, as
This Palaearctic species is adventive in North America where it is known from southern Newfoundland, Nova Scotia, New Brunswick, and Maine (Lindroth 1966: 631). The first inventoried specimen collected on this continent was found in Newfoundland before 1835. The Canadian Museum of Nature in Gatineau holds color drawings made by Philip Henry Gosse executed prior to 1834 of insects from Newfoundland and one drawing represents this species; the plates are bound together under the title
Four species, two endemic to northern Africa, one to Europe and western Asia, and one to North America.
The North American species is covered in Lindroth’s (1966: 633-634, as
Jeanne (1988: 77-78) pointed out the morphological differences between the type species of
This species ranges from Cape Breton Island (Bousquet 1987a: 126) to the Okanagan river in south-central British Columbia (Lindroth 1966: 634), south to western Oregon (Hatch 1953: 138), northeastern Kansas (Popenoe 1878: 78), and northwestern South Carolina (Anderson County, CNC). The record from “Georgia” (Bousquet and Larochelle 1993: 253) needs confirmation.
About 130 species (Liebherr and Schmidt 2004: 202-203) in the Nearctic (72 species, one adventive), Neotropical (15 species in Middle America, all but two shared with North America), Palaearctic (63 species, of which seven are Holarctic), and Afrotropical (three endemic species) Regions. The species are arrayed in four subgenera, following Liebherr and Schmidt (2004):
Liebherr (1994) published a key to all species found in the Western Hemisphere as well as a review of the Middle American species. Lindroth (1966, 1969a) covered all North American species except
Two species in North America, one of them is Holarctic.
The range of this species extends from Saskatchewan to Vancouver Island (Lindroth 1966: 589), north to southeastern Alaska (Lindroth 1966: 589), south to southeastern California (San Bernardino County, CNC; Casey 1920: 65) and southwestern Colorado (Casey 1920: 65, as
This Holarctic species is known from the Kamchatka Peninsula (Lindroth 1966: 584) and from Alaska (Lindroth 1966: 584) to Newfoundland (Lindroth 1955a: 129), south to northern New Hampshire (Coos County, Donald S. Chandler pers. comm. 2008), southeastern Michigan (Oakland County, UMAA), “Iowa” (Lindroth 1966: 584), east-central South Dakota (Brookings County, MCZ), east-central Colorado (Kiowa County, Foster F. Purrington pers. comm. 2010), “Idaho” (Lindroth 1966: 584), and northern Washington (Okanogan County, UASM). The record from “Pennsylvania” (Bousquet and Larochelle 1993: 255) needs confirmation.
Twenty-nine species in the Nearctic (16 species) and Palaearctic (17 species, six of them endemic to eastern Asia) Regions. Four species are Holarctic.
This species ranges along the west coast from southern British Columbia, including Vancouver Island (Lindroth 1966: 589, as
This species ranges from western Newfoundland (Lindroth 1955a: 125) to eastern Alaska (Lindroth 1966: 590), south to northeastern Washington (Gray 1937: 310, as
This species is found from Cape Breton Island (Bousquet 1987d: 106) to southern Manitoba (Goulet 1969: 280), south to east-central South Dakota (Larsen and Purrington 2010: 570), northern Illinois (Goulet 1969: 280), and northern West Virginia (Tucker and Preston Counties, CMNH). The record from “Nebraska” (Bousquet and Larochelle 1993: 253) needs confirmation.
A Holarctic species occurring in the Palaearctic Region from Norway to the Far East (Bousquet 2003c: 452) and in North America from Alaska (Lindroth 1966: 573) to Newfoundland (Lindroth 1955a: 126), south to Cape Breton Island (Lindroth 1954c: 307), Vancouver Island (Lindroth 1966: 573), and northern Washington (Glesne et al. 2000: 89). Fossil remnants, dated between 14,000 and 15,400 years B.P., have been unearthed in central Iowa (Schwert 1992: 78).
This species is restricted to a small area from Nova Scotia to the Ontario Peninsula (Lindroth 1966: 583), south to central Pennsylvania (Clinton County, CMNH) and New Jersey (Lindroth 1966: 583).
A Holarctic species ranging from the Kola Peninsula to eastern Siberia and in North America from Alaska to the Hudson Bay in Nunavut and northern Manitoba (Lindroth 1966: 574). Fossil remnants, dated between 14,000 and 20,700 years B.P., have been unearthed in east-central Iowa and central Illinois (Schwert 1992: 78).
This rarely collected species is known from Pennsylvania (Carbon, Chester and Greene Counties, Robert L. Davidson pers. comm. 2012), southernmost Ontario (Lindroth 1966: 583) and southern Michigan (Ingham County, CMNH) south to southeastern Texas (Casey 1920: 126) and eastern South Carolina (Kirk 1969: 12; Ciegler 2000: 112).
This widely distributed species ranges from western Newfoundland (Lindroth 1955a: 128) to Kodiak Island, north to the Anderson River delta in northern Northwest Territories (Lindroth 1966: 578), south to the Sierra Nevada in California (El Dorado County, CAS), San Miguel County in New Mexico (Fall and Cockerell 1907: 159, as
The range of this species extends from Cape Breton Island (Lindroth 1954c: 307) to Vancouver Island (Lindroth 1966: 581), north to southern Northwest Territories (Bousquet 1987a: 125), south to eastern Washington (Hatch 1953: 145), central Kansas (Trego and Scott Counties, CNC, CMNH), and northwestern Maryland (Garrett County, CMNH).
This species is found from southwestern New Brunswick (Webster and Bousquet 2008: 22) to Minnesota (Epstein and Kulman 1990: 214; Sherburne County, CNC), south to southwestern Nebraska (Lincoln County, Foster F. Purrington pers. comm. 2010) and along the Appalachian Mountains to central Alabama (Lee County, CNC).
This species ranges from the Nova Scotia Peninsula to south-central British Columbia, north to northern Alberta and Anticosti Island in Quebec, south to southeastern Wisconsin (Messer 2010: 42), the Saginaw Bay area in northern Michigan, and southern Vermont (Lindroth 1966: 574). Fossil remnants, dated about 10,800 years B.P., have been unearthed in central North Dakota (Ashworth and Schwert 1992: 260).
This species ranges from Newfoundland (Lindroth 1955a: 127) to western British Columbia (Lindroth 1966: 576), north to southern Northwest Territories (CNC), south to “Montana” (Casey 1920: 129, as
This rarely collected species is known from scattered localities from north-central Manitoba to the Gulf Coast of Alaska (Lindroth 1966: 571).
This species ranges from Newfoundland (Lindroth 1955a: 127) to central Alaska (Lindroth 1966: 576), south to “Oregon” (Hatch 1953: 144), southern Colorado (Wickham 1902: 239; LeConte 1879d: 500), northern Illinois (Lake County, CNC), and Maryland (Erwin 1981b: 154). The records from “Iowa” (Jaques and Redlinger 1946: 295), northeastern Kansas (Popenoe 1878: 78), and “North Carolina” (Bousquet and Larochelle 1993: 254) need confirmation.
This species ranges from New Brunswick (Bousquet 1987a: 125) to the Rocky Mountains in Alberta and eastern Alaska (Lindroth 1966: 579), south to northwestern Wisconsin (Messer 2010: 42), Beaver Islands, Lake Michigan, in northern Michigan (Dunn 1987: 11), and New England (Lindroth 1966: 579).
A Holarctic species widely distributed in the Palaearctic Region from Ireland to the Far East, south to Spain and Italy (Bousquet 2003c: 453). In North America, the taxon ranges from Alaska (Lindroth 1966: 580) to Newfoundland (Lindroth 1955a: 129), south to the District of Columbia (Ulke 1902: 7, as
Twenty-nine species in the Nearctic (four species, of which one extends into southern Mexico and one is adventive), Palaearctic (24 species, of which one is Holarctic), and Afrotropical (three species in eastern Africa including Madagascar) Regions.
A Holarctic species ranging in the Palaearctic Region across Siberia up to the Ob River (Lindroth 1966: 586), south to Kazakhstan and Mongolia (Bousquet 2003c: 454). In North America, the species ranges from Alaska and northern British Columbia to northeastern Alberta (Lindroth 1966: 586). The records from Colorado (Wickham 1902: 238) and New Mexico (Fall and Cockerell 1907: 159) probably refer to
This species ranges from Newfoundland (Lindroth 1955a: 125) to Vancouver Island (Lindroth 1966: 588), north to southern Yukon Territory (Hyland River, UASM), south to west-central California (Lindroth 1966: 587), southwestern Arizona (Maricopa County, UASM), New Mexico (Fall and Cockerell 1907: 159; UASM), and mountains of New England (Lindroth 1966: 587). The record from “Connecticut” (Bousquet and Larochelle 1993: 255) was based on a misidentified specimen of
This Palaearctic species is adventive in North America where it is known from west-central Alberta (Emerald Lake, Gerald J. Hilchie pers. comm. 2009) and southern British Columbia, including Vancouver Island (Lindroth 1966: 593), to northwestern California (Humboldt County, James R. LaBonte pers. comm. 1992) in the west and from Newfoundland (Lindroth 1955a: 121) to northeastern Minnesota (Kamal J.K. Gandhi pers. comm. 2008), south to northeastern Iowa (Larsen and Purrington 2010: 570) and northeastern West Virginia (Tucker County, CMNH) in the east. The first inventoried specimen collected in the east was found in Newfoundland around 1840 (Lindroth 1955a: 122) and in the west in 1933 (Lindroth 1966: 593; Leech 1935: 122, as
This widely distributed species ranges from Newfoundland (Lindroth 1955a: 120; Larson and Langor 1982: 594) to western British Columbia (Lindroth 1966: 614), south to Inyo County in California, the Sierra de Miahuatlán in Oaxaca (Liebherr 1994: 32), and northern Georgia (Fattig 1949: 34).
Seventy-two species in the Nearctic (50 species), Neotropical (14 species in Middle America but only two endemic), and Palaearctic (21 species, one Holarctic) Regions.
The range of this species extends from southeastern New Hampshire (Rockingham County, Ross T. Bell pers. comm. 2008) to eastern Kansas (Douglas County, Robert L. Davidson pers. comm. 2009), including southernmost Ontario (Lindroth 1966: 615), south to eastern Oklahoma (Latimer County, UASM), northwestern Louisiana (Bossier Parish, CMNH) and the Florida Panhandle (Jackson County, CNC). The record from “Iowa” (Bousquet and Larochelle 1993: 255) needs confirmation.
This species is known from a few localities in the Pacific Northwest from the Queen Charlotte Islands (Kavanaugh 1992: 74) to northwestern Oregon (Clackamas County, James R. LaBonte pers. comm. 1992).
This species ranges from central Prince Edward Island (CNC) and New Brunswick (Kouchibouguac National Park, CNC) to central Alaska (Lindroth 1966: 598), south at least to east-central California (Mono County, CAS), southeastern Arizona (Dajoz 2007: 21; UASM), northeastern New Mexico (San Miguel County, UASM), South Dakota including the Black Hills (Kirk and Balsbaugh 1975: 25), and northern Michigan (Dunn 1985a: 8; Casey 1920: 108, as
Lindroth (1966: 597) treated
The range of this species extends from Cape Breton Island (Lindroth 1954c: 305) to Vancouver Island (LeConte 1869c: 370), north to west-central Northwest Territories (Bousquet 1987a: 125), south to the Sierra Nevada in California (Casey, 1920: 106, as
This species is known from southwestern Oregon (Jackson County, CNC) to southern California (Fall 1901a: 46), including the Sierra Nevada (Papp 1978: 163). The record from “Idaho” (Horn 1872c: 385) is probably in error.
This species is found west of the Rocky Mountains from “Washington” (Hatch 1953: 143) and western Idaho (Nez Perce County, CNC) to southern California (Fall 1901a: 46). The record from Baja California (Horn 1894: 309) needs confirmation since the species is not listed in Liebherr’s (1994) review of the Mexican
This species is endemic to the Sierra Nevada [see Liebherr 1984b: Fig. 6], ranging from Lassen County to Tulare County.
This species ranges from west-central California south through the Coast Ranges to Baja California Norte [see Liebherr 1984b: Fig. 6].
This Holarctic species is found in eastern Siberia (Bousquet 2003c: 451) and in North America from Alaska (Lindroth 1966: 595) to Labrador (Goose Bay, CNC). The record from “Newfoundland” (Bousquet and Larochelle 1993: 258) refers to Labrador. Fossil remnants of this species, dated between about 14,000 and 18,100 years B.P., have been unearthed in central and southeastern Iowa (Baker et al. 1986: 96; Schwert 1992: 78).
The range of this species extends from southern Saskatchewan to Vancouver Island (Lindroth 1966: 592, as
This species ranges from southeastern Arizona to the Isthmus of Tehuantepec [see Liebherr 1994: Fig. 29].
The range of this species extends from southeastern Arizona to the Rio Grande drainage in south-central Texas, north to southeastern Colorado (Michels et al. 2008; Las Animas County, Robert L. Davidson pers. comm. 2008), south to the Federal District of Mexico [see Liebherr 1994: Fig. 28].
This species ranges from the Saguenay River in Quebec (Larochelle 1975: 35) to southeastern British Columbia (Lindroth 1966: 616), south to “Oregon” (Hatch 1953: 143), New Mexico (LeConte 1879b: 56; UASM), “Texas” (LeConte 1858a: 28, as
This species ranges from western New Hampshire (Grafton County, Ross T. Bell pers. comm. 2008) to eastern Oklahoma (Foster F. Purrington pers. comm. 2012), including southernmost Ontario (Lindroth 1966: 624), south to eastern Mississippi (Noxubee County, CMNH), southern Alabama (Löding 1945: 19), and central South Carolina (Ciegler 2000: 112). The record from northern Wisconsin along Lake Superior (Wickham 1896c: 134) is likely in error.
This species is known from a few localities along the Coastal Plain from southern Georgia (Fattig 1949: 33) and the Florida Panhandle (Peck and Thomas 1998: 23) west to northeastern Mississippi (Snodgrass and Cross 1983: 13), north along the Mississippi Basin to southern Illinois (Gallatin, Alexander, and Jackson Counties, CMNH, CNC). The record from “North Carolina” (Bousquet and Larochelle 1993: 260) needs confirmation.
This species ranges from New Brunswick (Lindroth 1966: 623) to southern Wisconsin (Messer 2010: 42), south to eastern Oklahoma (Latimer County, UASM), west-central Arkansas (Logan County, CNC), northern Georgia (Fattig 1949: 34), and eastern South Carolina (Ciegler 2000: 111). The records from “Minnesota” and “Louisiana” (Bousquet and Larochelle 1993: 256) need confirmation.
This species is found from eastern Arizona and western New Mexico south through the Sierra Madre Occidental to the Mexican states of Guerrero, Puebla, and Vera Cruz [see Liebherr 1986: Fig. 37].
This widely distributed species ranges from southern Quebec to Vancouver Island, south to Baja California Norte, the Isthmus of Tehuantepec, and central Florida; also present on several islands of the West Indies [see Liebherr 1986: Figs 51-54].
This species is found only in southern Georgia, throughout Florida including the Keys, and on
Liebherr (1986: 140) reported the presence of a series of putative hybrids
This species ranges from Cape Breton Island (Bousquet 1987d: 106) to northwestern Montana, including southern Manitoba and southern Saskatchewan (Ronald R. Hooper pers. comm. 2007), south to south-central Arizona, Jalisco in Mexico, and northern Florida [see Liebherr 1986: Fig. 46].
This species ranges from southern Arizona to southeastern Texas, south to southern Coahuila in Mexico [see Liebherr 1986: Fig. 41].
This species ranges from south-central Arizona south to the Mexican states of Sonora and southern Sinaloa along the Gulf of California Coast [see Liebherr 1986: Fig. 41].
This taxon ranges from central Kansas south to Chiapas in Mexico, west to central Arizona [see Liebherr 1986: Fig. 43].
This species ranges from Newfoundland (Lindroth 1955a: 123, as
This western species ranges from western British Columbia (Lindroth 1966: 609), as far north as the Skeena river area and Queen Charlotte Islands (Kavanaugh 1992: 75), south to southern California (Fall 1901a: 46, as
This species ranges from Rhode Island (Sikes 2003: 8; CMNH) to northeastern Illinois (Cook County, CMNH), south to eastern Texas along the Gulf Coast (Snow 1906a: 141) and central Florida (Brevard and Highlands Counties, CNC). The records from “Wisconsin” and “Pennsylvania” (Bousquet and Larochelle 1993: 258) need confirmation.
This species ranges from the Saint Lawrence Valley in southern Quebec (Larochelle 1975: 33) to the Fraser Valley in British Columbia (Lindroth 1966: 603), south to northern California (Siskiyou County, CNC), east-central California (Mono County, Foster F. Purrington pers. comm. 2012), northern Arizona (Coconino and Apache Counties, CNC), southern Colorado (Alamosa County, CNC), and southern Nebraska (Chase County, Foster F. Purrington pers. comm. 2010). The record from “Michigan” (Bousquet and Larochelle 1993: 256) needs confirmation.
This rarely collected species is found along the provinces and states bordering the Atlantic Coast from Cape Breton Island (Lindroth 1966: 611) to Connecticut (New London County, William L. Krinsky pers. comm. 2008).
This species ranges from Cape Breton Island (Lindroth 1954c: 306) to western Wisconsin (Kleintjes et al. 2003: 81; Messer 2010: 42), south to northeastern Mississippi (Snodgrass and Cross 1983: 13) and west-central South Carolina (Ciegler 2000: 112). The record from “Prince Edward Island” (Bousquet and Larochelle 1993: 256) is based on misidentified small specimens of
This species ranges from Cape Breton Island (Lindroth 1954c: 306) to Vancouver Island (Lindroth 1966: 609), south to “Oregon” (CAS), north-central Idaho (Idaho County, CNC), southeastern Wyoming (Lavigne 1977: 43), southern Minnesota (Hennepin County, CNC), northeastern Illinois (Purrington et al. 2002: 200), and southeastern Virginia (Surry County, CMNH). The record from “Nebraska” (Bousquet and Larochelle 1993: 257) is probably in error.
This species is known only from the holotype which has no locality or state label (see Liebherr 1991a: 121-122).
None.
The range of this species extends from Cape Breton Island (Lindroth 1954c: 306) to southwestern British Columbia (Lindroth 1966: 600), south to northwestern California (Trinity County, CNC), northern Colorado (Haubold 1951: 705; Armin 1963: 150), west-central Kansas (Trego County, CNC), and North Carolina (Haywood County, CNC).
This species ranges from Newfoundland (Lindroth 1955a: 122) to the Queen Charlotte Islands (Kavanaugh 1992: 74), north to southern Northwest Territories (Bousquet 1987a: 125), south to northern Idaho (Peter W. Messer pers. comm. 2008), northwestern Montana (Glacier County, CNC), northern Wisconsin (Wickham 1896c: 133; Casey 1920: 114, as
This species is known from western Maine (Franklin, Kennebec, and York Counties, Ross T. Bell pers. comm. 2008), southwestern Quebec, the Ontario Peninsula, and southern Manitoba (Lindroth 1966: 611) south to southern Louisiana (Assumption and East Baton Rouge Parishes, LSAM) and northern Florida (Alachua County, CNC).
This species is found from Newfoundland (Lindroth 1955a: 124) to British Columbia, north to east-central Alaska (Lindroth 1966: 601), south to west-central Washington (Lindroth 1966: 601), northwestern Montana (Russell 1968: 64), central Indiana (Downie 1957: 116; Schrock 1985: 343), and eastern South Carolina (Kirk 1969: 11; Ciegler 2000: 112).
This widely distributed species extends from Newfoundland (Lindroth 1955a: 124) to Alaska (Lindroth 1966: 612), south to the Sierra Nevada in northeastern California (Liebherr 1991a: 119), southwestern Colorado (Wickham 1902: 239; Ouray County, CNC), northwestern Nebraska (Sheridan County, CNC), northwestern Pennsylvania (Erie County, CMNH), and “Maryland” (Liebherr 1994: 30). One specimen is also known from the city of Mexico (Liebherr 1994: 30) but in my opinion it is likely mislabeled.
This species ranges from Cape Breton Island (Lindroth 1954c: 306) to northeastern Minnesota (Kamal J.K. Gandhi pers. comm. 2008), south to southern Georgia (Fattig 1949: 34). The record from “Nebraska” (Bousquet and Larochelle 1993: 258) needs confirmation.
This species is known from Cape Breton Island (CNC) to southern Manitoba (Lindroth 1966: 601), south to southwestern North Carolina (Macon County, CNC).
This species ranges from Nova Scotia (Lindroth 1954c: 306) to southeastern South Dakota (Kirk and Balsbaugh 1975: 25), south at least to northeastern Kansas (Popenoe 1877: 23), central Louisiana (East Feliciana Parish, CMNH, CNC), and the Florida Panhandle (Peck and Thomas 1998: 22).
This species ranges from “New York” (LeConte 1854b: 52, as
This species ranges from central North Carolina (Casey 1920: 86, as
This eastern species is found from southern Quebec (Larochelle 1975: 38) to eastern South Dakota (Kirk and Balsbaugh 1975: 24), south to Oklahoma (French et al. 2001: 228; Latimer County, UASM) and the District of Columbia (Ulke 1902: 7). The records from central Arizona (Griffith 1900: 565), northern Colorado (Wickham 1902: 239; Armin 1963: 149), southeastern Texas (Snow 1906a: 141), “Mississippi,” “Alabama” (Bousquet and Larochelle 1993: 259), and “Florida” (LeConte 1879b: 56) are probably in error; those from “Maryland,” “Delaware” (Bousquet and Larochelle 1993: 259), and South Carolina (Kirk 1970: 12, as
This species ranges from New Jersey (Essex County, CMNH) to east-central Iowa (Muscatine County, Doug A. Veal pers. comm. 2010), including southernmost Ontario (Lindroth 1966: 619), south to southeastern Texas (Casey 1920: 88) and northern Florida (Peck and Thomas 1998: 23). The record from “Wisconsin” (Bousquet and Larochelle 1993: 259) needs confirmation.
This species ranges from Nova Scotia (Lindroth 1966: 594) to southeastern Manitoba (UASM), south to eastern Texas (Orange and Sabine Counties, CNC; Dajoz 2007: 23) and central Florida (Peck and Thomas 1998: 23).
The range of this species extends from Cape Breton Island (Bousquet 1987a: 125) to southeastern South Dakota (Kirk and Balsbaugh 1975: 25), south to southeastern Nebraska (Hall County, Foster F. Purrington pers. comm. 2009), east-central Missouri (Saint Louis, CNC), and northwestern South Carolina (Kirk 1970: 12; Ciegler 2000: 111). The records from southern Louisiana (Summers 1874a: 80), “Kansas,” and “Arkansas” (Bousquet and Larochelle 1993: 259) need confirmation.
This species ranges from “Maryland” (CMNH) to north-central Oklahoma (Kay County, CNC), south at least to central Texas (Travis and Blanco Counties, CMNH; Casey 1920: 73), eastern Louisiana (Washington Parish, LSAM), and the Florida Panhandle (Peck and Thomas 1998: 23). The record from “Kansas” (Bousquet and Larochelle 1993: 259) needs confirmation.
This species ranges from southeastern New Hampshire (Rockingham County, Ross T. Bell pers. comm. 2008) to southeastern Colorado (Michels et al. 2008), including southern Ontario (CNC), south to southern Mexico (Liebherr 1994: 20) and southern Florida (Peck and Thomas 1998: 23), west along the southwest to southern California (Liebherr 1994: 19-20); also recorded from the Bahamas and Bermuda where it is probably adventive (Hilburn and Gordon 1989: 677; Liebherr 1994: 19). The record from Quebec (Lindroth 1966: 622) refers to
This species ranges from New Jersey (Ocean County, CMNH) south to northwestern Louisiana (Bossier Parish, LSAM), southwestern Mississippi (Hancock and Hinds Counties, CMNH), southwestern Alabama (Casey 1920: 75, as
The range of this eastern species extends from southeastern New Hampshire (Rockingham County, Donald S. Chandler pers. comm. 2009) to eastern South Dakota (Kirk and Balsbaugh 1975: 25), including southern Ontario (Lindroth 1969a: 1120), south to southeastern Texas (Casey 1920: 74) and southeastern Georgia (Fattig 1949: 32, as
This species is known from scattered localities from southwestern Indiana (Blatchley 1910: 123) south to southwestern Georgia (Fattig 1949: 34), southwestern Alabama (Casey 1920: 71; Mobile County, MCZ), southeastern Mississippi (George County, MCZ), north-central Louisiana (Winn Parish, MCZ), and northeastern Texas (Wood County, UASM). The record from Alberta (see Lindroth 1966: 631) must be in error after all; those from northwestern New York (Notman 1928: 235) and “Michigan” (Bousquet and Larochelle 1993: 261) need confirmation.
The number of species of
Liebherr and Will (1996: 303-307) published a key for the identification of all species found in North America. Bousquet (2012: 77-78) described a new species and issued a key for the species found east of the Mississippi River.
Two species in southwestern North America.
This species is restricted to southern California from the Santa Ynez Mountains to San Diego County (Liebherr and Will 1996: 318).
This species is known only from the Sierra Blanca (Barr 1982b: 98), Capitan, and Sacramento Mountains (Liebherr and Will 1996: 318) in New Mexico.
Sixteen species in the Nearctic (nine species, of which one extends into Mexico) and Palaearctic (seven species) Regions.
This species ranges from southern British Columbia, including Vancouver Island (Lindroth 1966: 639), south to the northern parts of the Baja California Peninsula (Horn 1894: 309), east to western New Mexico (Liebherr and Will 1996: 316) and northern Sonora (Bates 1884: 280).
This species occurs along the Appalachian Mountains from Connecticut to southern Pennsylvania, south to northwestern Alabama [see Bousquet 2012: Fig. 4]
This species ranges from Newfoundland (Lindroth 1955a: 118, as
This species ranges from Nova Scotia (Majka et al. 2007: 11) to eastern Ohio, south to West Virginia and Maryland [see Liebherr and Will 1996: Fig. 18].
This species ranges from southwestern New Brunswick (Webster and Bousquet 2008: 22) to northeastern Iowa (Allamakee County, Doug A. Veal pers. comm. 2009), including the upper peninsula of Michigan (Casey 1920: 30, as
This species is found along the west coast from Vancouver Island (Lindroth 1966: 645) to at least west-central California (Marin County, CNC). The record from northern Colorado (Wickham 1902: 238; Armin 1963: 153) is in error.
This species ranges from southwestern Pennsylvania southwestwards to eastern Oklahoma [see Bousquet 2012: Fig. 4].
The range of this species extends from Cape Breton Island (Lindroth 1966: 643) to the Black Hills in southwestern South Dakota (Kirk and Balsbaugh 1975: 24), south to northern Colorado (Armin 1963: 154; UASM), northwestern Texas (Wheeler County, CMNH), northeastern Oklahoma (Delaware County, CMNH), and southern South Carolina (Ciegler 2000: 115). The record from “Florida” (Bousquet and Larochelle 1993: 262) needs confirmation.
This species is known from high mountains along the Tennessee - North Carolina border (Barr 1969: 81); it was also reported from northeastern Georgia (Fattig 1949: 33).
Forty-four species are recorded by Lorenz (2005: 429-430), five in the Nearctic Region, with one of them Holarctic, and 40 in the Palaearctic Region. A number of Palaearctic species considered as
This species is found from the Adirondack Mountains in New York (Casey 1920: 9 as
This species ranges from Sable Island (CNC) off the Nova Scotia coast to southeastern Minnesota (Gandhi et al. 2005: 932), south to southern Texas (Casey 1920: 40, as
This species is found from Vermont (Rutland County, CNC) to the lower peninsula of Michigan (Hubbard and Schwarz 1878: 644), including southern Ontario (Lindroth 1966: 646), south to east-central Kentucky (Powell County, CNC). The record from “Iowa” (Bousquet and Larochelle 1993: 263) needs confirmation; that from “Virginia” (Bousquet and Larochelle 1993: 263) is probably in error (Richard L. Hoffman pers. comm. 2009).
Morvan (2000: 76) treated this taxon as a subspecies of
This Holarctic species ranges from Newfoundland (Lindroth 1955a: 119) to central British Columbia and Alaska (Lindroth 1966: 636), south to northeastern Minnesota (Casey 1920: 36, as
This characteristic species is known for sure only from the holotype. However, Fattig (1949) recorded it also from Dalton, Georgia but the specimen(s) was apparently destroyed (Liebherr 1990: 432).
Thirty-six species in Middle America (28 species, of which one extends into southern North America and two into South America) and the West Indies (eight species).
Liebherr (1992) revised the species.
This species is known from southeastern Arizona and the northern Sierra Madre Occidental in Sonora and Chihuahua [see Liebherr 1992: Fig. 160].
Four species are assigned to this subgenus, one from southwestern United States and three from Mexico (Liebherr and Will 1996: 316).
This species is known from several localities in southeastern Arizona [see Liebherr and Will 1996: Fig. 24].
About 320 species are listed in this subgenus by Lorenz (2005: 430-433). Of these only five unrelated species are found in southern United States, two of them being endemic (
This species is known only from Pima and Santa Cruz Counties in southeastern Arizona and from one specimen without precise locality collected in Mexico [see Liebherr and Will 1996: Fig. 24].
This species is known from Pima, Cochise, and Santa Cruz Counties in southern Arizona (Liebherr and Will 1996: 318).
Whitehead (1973: 196) stated that adults of this form are possibly conspecific with those of
This species ranges from southern Utah, eastern Arizona, and western New Mexico south to Oaxaca in Mexico; it also occurs throughout the Baja California Peninsula (Liebherr and Will 1996: 318).
The range of this species extends from southern Arizona and western Texas (Jeff Davis County, CMNH) south to the state of Oaxaca (Whitehead 1973: 201).
This species is yet known only from Cochise, Pima, and Santa Cruz Counties in southern Arizona (Liebherr and Will 1996: 318). The record from southwestern Utah (Tanner 1928: 270) is probably in error.
Twenty-three species in the Palaearctic (11 Asian species) and Oriental (15 species) Regions are listed in this genus by Lorenz (2005: 423). Liebherr (2005), on the other hand, redefined cladistically the genus to include seven Pacific and Asian species. One of these species is adventive in western North America.
The sole species present in North America is included in Hatch’s (1953: 132) monograph on the beetles of the Pacific Northwest. It is also described in length by Habu (1978: 126-128).
This adventive species is known in North America from a few specimens collected in southern British Columbia (Jarrett and Scudder 2001: 382), southwestern Washington (Pacific County, Peter W. Messer pers. comm. 2011), western Oregon (Hatch 1953: 132), and northern Idaho (Liebherr and Will 1996: 303). The first specimen collected on this continent was found in 1931 in Portland, Oregon (Hatch 1953: 132). The species is also adventive in Hawaii (Liebherr and Zimmerman 2000: 466).
About 120 species (Lorenz 2005: 438-439), mainly in the tropical regions, arrayed in five genera. Only one endemic species (
Excluding the subcosmopolitan
Eighteen species in the Nearctic (one endemic species), Australian (seven endemic species), and Afrotropical (ten endemic species) Regions and one (
Bousquet (1987a: 128) commented on the structural differences between the two species found in North America.
The relationship of the endemic North American species (
This species is subcosmopolitan. It is adventive in North America where it is known from Nova Scotia (Pictou County, CNC) to eastern Iowa (Muscatine County, Doug A. Veal pers. comm. 2009), south to east-central Texas (Riley 2011), central Louisiana (Grant Parish, CNC) and southern Florida (Peck and Thomas 1998: 23) in the east, and from Oregon (Hatch 1953: 147) to southern California (Fall 1901a: 47) in the west. The first inventoried specimen collected on this continent was found prior to 1853 (LeConte 1853c: 386, as
The range of this species extends from southwestern Maine (Nelson 1991: 284) to eastern Iowa (Wickham 1911b: 7), including southernmost Ontario, south to “Missouri,” “Alabama” (Bousquet 1987a: 127-128), and northern Florida (Hamilton, Okaloosa, and Taylor Counties, CNC).
One genus with two species in the Nearctic and Palaearctic Regions.
Two species in the temperate regions of eastern North America (
Lindroth (1968: 649) commented on the structural differences between the two species.
The range of this species extends from central Nova Scotia (Hants County, Christopher G. Majka pers. comm. 2008) to southwestern Nebraska (Foster F. Purrington pers. comm. 2010), south to “Texas” (Barr 1964: 3; Lindroth 1968: 649) and west-central South Carolina (Ciegler 2000: 108).
About 115 species (Lorenz 2005: 440-441, as
Ball and Bousquet (2000: 108) recognized two groups of genera within this tribe. The eucaerine complex comprising
Western Hemisphere, with 26 species (Lorenz 2005: 440) in the Nearctic (two species) and Neotropical (24 species) Regions.
Schaeffer (1910: 395) commented on the structural differences between the two species found in North America.
This species is known only from Texas (Val Verde County, CNC).
This species is known from southeastern Arizona (Graham County, CNC) and southern Texas (Schaeffer 1910: 395).
Western Hemisphere, with about 35 species in temperate, subtropical, and tropical areas of the Nearctic (one species only) and Neotropical Regions.
According to Liebherr (1988: 34),
This species ranges from Kansas (LeConte 1858a: 28; Horn 1872c: 385) to southwestern Oregon (Hatch 1953: 146; Jackson County, MCZ, UASM), south to southern California (Moore 1937: 11; San Diego County, CNC, MCZ) and Costa Rica (Liebke 1936: 463).
Four species in temperate, subtropical, and tropical areas of the Nearctic (two species) and Neotropical (three species) Regions.
Horn (1891: 38) commented on the structural differences between the two species found in North America.
This species is known so far only from southwestern California (Horn 1891: 38; Fall 1901a: 47; Moore 1937: 11).
This species is found along the Coastal Plain from southeastern North Carolina to central Florida, west to northeastern Texas (Dallas County, MCZ) and central Oklahoma (Grady County, Robert L. Davidson pers. comm. 2012), including west-central Louisiana (Allen 1965: 67), along the Mississippi Basin in southwestern Ohio (Blatchley 1910: 137) and central Indiana (Downie and White 1967: 308), in the West Indies from the Bahamas (Turnbow and Thomas 2008: 12) to Trinidad, in tropical mainland from central Mexico to Santa
Twenty-five species (Lorenz 2005: 440-441) in the Western Hemisphere arrayed in two subgenera:
Seventeen species in the temperate, subtropical, and tropical areas of the Nearctic (two species) and Neotropical (17 species) Regions.
Horn (1881: 153) commented on the structural differences between the two species found in North America.
This species ranges from southernmost Nevada (Clark County, CNC) south to Guatemala (Bates 1883a: 157), including southeastern California (Riverside County, MCZ, UASM) and southern Arizona (Fall 1901a: 48; Maricopa and Pima Counties, MCZ, UASM). The record from “Colorado” (LeConte 1858a: 28) probably refers to the Colorado River.
This species is known along the Coastal Plain from South Carolina (Kirk 1969: 12; Ciegler 2000: 116; Clarendon County, MCZ) to southern Florida (Peck and Thomas 1998: 23), west to central and southeastern Texas (Lee and Cameron Counties, MCZ; Wickham 1897: 107), and south to Nicaragua (Bates 1883a: 157). The record from “Arizona” (Ball and Bousquet 2000: 109) refers to
Fourteen species in the Western Hemisphere arrayed in two subgenera:
Eleven species in temperate, subtropical, and tropical areas of the Nearctic (one species) and Neotropical (ten species) Regions arrayed in two species groups (Ball and Hilchie 1983: 107-108).
This species inhabits the Coastal Plain from central Maryland (Hoffman et al. 2006: 28) to southern Florida (Peck and Thomas 1998: 23), west to eastern Texas (Hardin County, UASM), including southeastern Mississippi (Jackson County, Drew A. Hildebrandt pers. comm. 2008).
Worldwide but excluding Europe and northern Africa, with about 165 species (Lorenz 2005: 445-447) arrayed in five genera:
About 85 species in the temperate, subtropical, and tropical areas of the Nearctic (six species, only one endemic), Neotropical (about 25 species), Australian (10 species), Oriental (about 20 species), Palaearctic (11 Asian species), and Afrotropical (25 species) Regions.
Lindroth’s (1969a: 1013) key covered all North American known at the time. Two new species have been described subsequently by Bell (1987) and Mateu (1995).
This species ranges from Colorado County in southeastern Texas to Guatemala (Reichardt 1968: 150).
This species is known from a few localities from southeastern Arizona and southwestern New Mexico (Bell 1987: 373) south to Chiapas in southern Mexico (Bell 1989a: 156).
This subspecies ranges from southwestern New Jersey (Smith 1910: 212) and northern Virginia (Hoffman and Roble 2000: 39) to northeastern Kansas (Popenoe 1877: 23, as
The subspecies
This species is known from southeastern Georgia (Camden and Glynn Counties, CMNH), central Florida (Pinellas County, CMNH), north-central Mississippi (Grenada County, Drew A. Hildebrandt pers. comm. 2010), and east-central (Riley 2011) and south-central Texas (Bastrop County, Peter W. Messer pers. comm. 2010).
This species is known only from the Florida Peninsula as far north as Alachua County (Peck and Thomas 1998: 22), the Bahamas (Turnbow and Thomas 2008: 14), Cuba (Bell 1985a: 323), and the Cayman Islands (Darlington 1947: 211).
This species ranges from southern Quebec (Larochelle 1975: 98) to western Wisconsin (Messer 2010: 42), south to Guatemala (Bates 1883a: 217), southeastern Louisiana (Saint Tammany Parish, Igor M. Sokolov pers. comm. 2009), and central Florida (Vince Golia pers. comm. 2007), west to southern Arizona and the Baja California Peninsula (Horn 1894: 311).
Worldwide, with about 325 species (Lorenz 2005: 441-445, as
Western Hemisphere, with about 100 species (Lorenz 2005: 442-444, as
Bousquet (2010b) reviewed the Nearctic, Mexican and West Indian (Greater Antilles) species and provided a key for their identification.
Liebke (1930, 1938) recognized an array of subgenera and several appear to be polyphyletic assemblages. A taxonomic revision of the genus is much needed. Lorenz (2005: 442) recognized
Three species in the Neotropical Region (Liebke 1938: 57), one of them extending into southern Arizona.
This species ranges from southern Arizona (Bousquet 2010b: 7) to Panama (FSCA).
Fourteen species are listed in this subgenus by Lorenz (2005: 442) but some (e.g.,
This species has a very restricted distribution; it is known from southeastern Arizona and adjacent northern Mexico [see Bousquet 2010b: Fig. 19].
This species ranges from Maine (Dearborn and Donahue 1993: 8; Foss 2001: 14) to eastern South Dakota (Kirk and Balsbaugh 1975: 37), including southern Quebec (Larochelle 1975: 78) and the Ontario Peninsula (Lindroth 1969a: 1008), south to the Yucatán Peninsula and southern Florida, west to southern Arizona and Baja California Sur [see Bousquet 2010b: Fig. 18]; also known from Cuba and Jamaica (Bousquet 2010b: 10).
Five Nearctic and Neotropical species were included in this subgenus by Liebke (1938: 55-56). However, a number of species (e.g.,
This species is known from southern Florida, Cuba, the Cayman Islands, Jamaica, and Haiti (Bousquet 2010b: 23).
This species ranges from southeastern Arizona (Bousquet and Larochelle 1993: 334) south at least to Honduras (USNM).
This species inhabits the Coastal Plain and ranges from southeastern New York to southern Florida, west to eastern Louisiana [see Bousquet 2010b: Fig. 20]. The species was also recorded from “N. Yucatan” by Liebke (1938: 56) and from “Pennsylvania” by Leng (1920: 65); both records are suspect.
Western Hemisphere, with about 65 species (Lorenz 2005: 393) arrayed in 16 genera.
Thirty-four species (Lorenz 2005: 393) in the temperate, subtropical, and tropical areas of the Nearctic (three species) and Neotropical (32 species) Regions.
Bousquet (1997b: 336) commented on the structural differences between
This species is known only from the Florida Peninsula (Peck and Thomas 1998: 23).
This species ranges from Rhode Island (William L. Krinsky pers. comm. 2009) and Connecticut (Krinsky and Oliver 2001: 232) to southeastern South Dakota (Kirk and Balsbaugh 1975: 36), including the Ontario Peninsula (Lindroth 1969a: 1006), south to southeastern Texas (Wharton County, CNC) and southern Florida (Peck and Thomas 1998: 23); also found in Cuba (Gundlach 1891: 19), Barbados, and Hispaniola (Peck and Thomas 1998: 23).
This species is known from a few scattered localities in Tennessee, Louisiana (Bousquet 1997b: 338), and Texas (Chaudoir 1872d: 412).
About 120 species in the Nearctic (four species), Neotropical (about 40 species of
This tribe is treated here in a restricted sense following Ball and Bousquet (2000: 109) and excluded the graphipterines, corsyrines, masoreines, somoplatines, and nemotarsines.
About 75 species (Lorenz 2005: 453) arrayed in two subgenera,
Lindroth (1969a: 1010) included a key to four of the five species found in North America.
Western Hemisphere, with about 40 species in the Nearctic (five species, one of them adventive) and Neotropical (about 40 species) Regions. One South American species (
This species ranges from southwestern Maine (Majka et al. 2011: 47) and southern Quebec (CNC) to southwestern Minnesota (Gandhi et al. 2005: 932), south to southern Texas (Wickham 1897: 107; Zapata County, CMNH) and central Florida (Peck and Thomas 1998: 23), west along the south to Riverside County in California (Dajoz 2007: 19) and the Baja California Peninsula (Horn 1894: 310; Horn 1895: 226).
This species ranges from North Carolina (Moore County, CNC) to southern Arizona (Pima County, CMNH), south to Jalisco in Mexico (CMNH), southern Texas (Johnson 1978: 68), and southern Florida (Peck and Thomas 1998: 23); also known from the Bahamas (Darlington 1953: 16). The record from “Virginia” (Bousquet and Larochelle 1993: 266) needs confirmation; that from southern Ontario (Belaoussoff et al. 2003: 878) probably refers to
This South American species is adventive in southeastern Florida (Shpeley and Ball 2008: 9). The first inventoried specimen collected on this continent was found in March 2007 (Halbert 2007: 7; Shpeley and Ball 2008: 9). In South America, the species ranges from the Baía de Marajó along the Atlantic Coast in northern Brazil to southern Uruguay and central Argentina (Shpeley and Ball 2008: Fig. 6).
This species occurs from southwestern Alabama (Baldwin and Clarke Counties, CMNH) to central Arizona (Yavapai County, CNC), south to southern Texas (Wickham 1897: 107; Johnson 1978: 68); also recorded from Mexico (Blackwelder 1944: 52).
This species is known from southwestern California (Fall 1901a: 48; Moore 1937: 11) to the western tip of Texas (El Paso County, CMNH), including western Nevada (Bechtel et al. 1983: 474); also recorded from the Baja California Peninsula (Horn 1895: 226).
Worldwide, with about 4,260 species arrayed in 237 genera (Lorenz 2005: 453-502, as
Worldwide, with about 825 species arrayed in 77 genera (Lorenz 2005: 454-464). The subtribe is underrepresented in the Northern Hemisphere with about 70 species (8.5% of the world fauna), of which nine (about 1%) only occur in North America.
Shpeley and Ball (2001) reviewed all 111 native species-group taxa found in the Western Hemisphere and provided keys for their identification. Their work is to be used in conjunction with a previous study (Ball and Shpeley 1983).
In a cladistic analysis based on characters of the adults performed by Ball et al. (1995: Fig. 7), this subtribe is positioned as the sister-group to {
Nine species (Lorenz 2005: 460) in the Palaearctic (eastern Asia only), Oriental, and Australian Regions, with one of them adventive in Florida.
The character states of the sole species found in North America are described in detail by Habu (1967: 104-105) under the name
Habu (1967: 100) treated this taxon as a subgenus of
This species is widely distributed in Asia from India to Japan, south through the Malay Archipelago to New Guinea; it is also known from Christmas Island and Samoa. The species is adventive in southeastern United States and currently known from Florida, where the first specimens were detected in 1992 in Palm Beach County (Choate 2001), southern Mississippi (Pearl River County, Peter W. Messer pers. comm. 2010), and east-central Louisiana (West Feliciana Parish, Igor M. Sokolov pers. comm. 2009).
Thirty-four species in the temperate, subtropical, and tropical areas of the Nearctic (one eastern species) and Neotropical (34 species) Regions. These species are arrayed in five subgenera:
Shpeley and Ball (2001: 78-80) published a key for the identification of all known species.
Nine species in North America and Middle America.
This species extends from Maryland (Steiner et al. 2007: 224) to central Florida (Peck and Thomas 1998: 24) and southeastern Louisiana, and from the states of Tamaulipas and Jalisco in Mexico to Panama [see Ball 1975: Fig. 112].
Twenty-four species (Lorenz 2005: 463-464) in the Neotropical Region, one of them reaching southeastern Texas. The species are arrayed in five subgenera:
Ten species in the Neotropical Region, one of them reaching southeastern Texas.
This species ranges from southern Texas (Shpeley and Ball 2001: 162) south to northeastern Brazil [see Ball and Shpeley 1983: Fig. 62].
Two species from the Afrotropical Region, one restricted to Madagascar, the other subcosmopolitan, having been dispersed through commerce.
The character states of the species found in North America are described in detail by Habu (1967: 75-77).
This genus was listed in the tribe
This species is adventive in North America where it is known from Randolph County in North Carolina (Peter W. Messer pers. comm. 2012), Aiken County in South Carolina (Peter W. Messer pers. comm. 2011), Telfair and Jeff Davis Counties in southern Georgia (Harry J. Lee, Jr. pers. comm. 2010), Broward County in Florida (Peck and Thomas 1998: 25), Polk and Garland Counties in Arkansas (CMNH), and Santa Cruz County in southern Arizona (Ober and Maddison 2008: 30, as
About 105 species (Lorenz 2005: 457-458) in the Nearctic (five species), Neotropical (43 species), Australian, Afrotropical, Oriental, and Palaearctic (13 species in eastern Asia only) Regions. These species are arrayed in four subgenera:
About 45 species in temperate, subtropical, and tropical areas of the Western Hemisphere arrayed in 11 species groups. Five species are found in North America.
Shpeley and Ball (1994) revised the species and provided a key for their identification.
The range of this species extends from Connecticut (New London County, William L. Krinsky pers. comm. 2012) to southeastern Nebraska, south to eastern Texas and west-central Florida [see Shpeley and Ball 1994: map 19]. According to Shpeley and Ball (1994: 119), the specimens labeled from Gallatin County in Montana (CAS) and Santa Cruz County in Arizona (AMNH) are probably mislabeled.
This species is found from southeastern Arizona and southeastern New Mexico south to Honduras and Belize [see Shpeley and Ball 1994: map 18].
The range of this species extends from southern Arizona and southwestern New Mexico south to southeastern Brazil and northern Argentina [see Shpeley and Ball 1994: map 17].
This species occurs in southern Florida, the West Indies, and from Mexico (as far north as San Luis Potosí) to Brazil [see Shpeley and Ball 1994: map 12].
This species is known from the tip of Florida, the West Indies, including the Bahamas (Turnbow and Thomas 2008: 12), Cuba, and the Dominican Republic (Shpeley and Ball 2001: 148), and from the state of San Luis Potosí in Mexico to southeastern Brazil [see Shpeley and Ball 1994: map 16].
About 315 species (Lorenz 2005: 465-470, excluding
About 200 species (Lorenz 2005: 465-469, as
Ball and Hilchie (1983) regarded
Twenty-one species in North America (16 species, one of them Holarctic), Mexico (two species shared with North America), Eurasia (two species, one endemic to eastern Asia), and North Africa (four endemic species).
Lindroth (1969a) reviewed all North American species. Since then, one Palaearctic species (
Lorenz (2005) listed all Nearctic species in the subgenus
This species ranges from southern Quebec (Larochelle 1975: 78) to eastern South Dakota (Kirk and Balsbaugh 1975: 39), south to northeastern Texas (Lamar County, Brian Raber pers. comm. 2010), southern Louisiana (Chaudoir 1873c: 103), northern Alabama (Madison County, CMNH), and southeastern South Carolina (Ciegler 2000: 119). The record from “Florida” (Bousquet and Larochelle 1993: 267) needs confirmation.
This species ranges from southwestern California to “New Mexico” (Lindroth 1969a: 1075).
The range of this species extends from Newfoundland (Lindroth 1955a: 132) to southern British Columbia, south to “New Mexico” (Lindroth 1969a: 1082), northern Wisconsin along Lake Superior (Wickham 1896c: 135; MCZ), northwestern Pennsylvania (Erie County, CMNH), and Connecticut (Krinsky and Oliver 2001: 272). The record from “South Dakota” (Bousquet and Larochelle 1993: 267) is probably in error.
This species is known so far only from the holotype (Lindroth 1969a: 1086) collected in southwestern California.
This widely distributed species ranges from Newfoundland (Lindroth 1955a: 131) to Vancouver Island, north to Yukon Territory (Lindroth 1969a: 1076), south to “Oregon” (Horn 1882: 152), southeastern Arizona (Snow 1906b: 162), southern New Mexico (Otero County, CMNH; Lindroth 1969a: 1076), northeastern South Dakota (Kirk and Balsbaugh 1975: 38), and western Maryland (Bailey et al. 1994: 320). The records from “Prince Edward Island” (Bousquet and Larochelle 1993: 267, see Majka et al. 2008: 133), “Kansas” (Horn 1872c: 385), and “Nebraska” (Wickham 1896c: 135) need confirmation; that from southern California (Moore 1937: 12) is probably in error though one old specimen labeled “Cal.” is known (MCZ).
This species is found from Massachusetts (Darlington 1936b: 148; Hampden County, MCZ) south to the Florida Panhandle (Peck and Thomas 1998: 24) and southwestern Alabama (Casey 1920: 288).
This species is known from a few localities in southern Oregon (Westcott et al. 2006: 8; Harney County, CMNH; Lake County, Foster F. Purrington pers. comm. 2009), west-central Idaho (Valley County, USNM), southwestern Wyoming (Parmenter and MacMahon 1984: 26), south-central Utah (Casey 1913: 179; Knowlton 1939: 2; Piute County, MCZ), “Nevada” (CMNH, MCZ), and east-central California (Lindroth 1969a: 1079). The record from Seattle, Washington (Hatch 1953: 158) needs confirmation.
This species is found from the southern part of the Prairie Provinces south to southeastern Arizona (Graham County, UASM), central New Mexico (Lindroth 1969a: 1074), western Texas (Ward County, Ken Karns pers. comm. 2009), and northwestern Oklahoma (Woods County, CMNH).
This species ranges from northwestern South Dakota (Kirk and Balsbaugh 1975: 38) to western Montana (Hatch 1933a: 9), south to northern Sonora (Bates 1884: 296) and southern Texas (Sutton, Val Verde, Edwards, and Colorado Counties, CMNH). The record from “Arkansas” (Bousquet and Larochelle 1993: 268) needs confirmation; those from southern Wisconsin (Rauterberg 1885: 14) and southwestern California (Moore 1937: 12) are probably in error.
The range of this species extends from Sable Island (Christopher G. Majka pers. comm. 2007) off the coast of Nova Scotia to southern Alberta (Lindroth 1969a: 1086), south to northwestern South Dakota (Kirk and Balsbaugh 1975: 39) and southern South Carolina (Ciegler 2000: 119).
This species ranges from southern Quebec (Larochelle 1975: 79) to southeastern North Dakota (Ransom County, CNC), north to southwestern Manitoba (Stjernberg 2011: 71), south to “Texas” (Lindroth 1969a: 1083) and northeastern Florida (Duval County, USNM). The record from the state of Puebla in Mexico (Bates 1891a: 270) needs confirmation.
The range of this species extends from southern Manitoba to the Okanagan Valley in south-central British Columbia, north to southern Northwest Territories (Lindroth 1969a: 1078), south to northern Oregon (Baker and Wasco Counties, CMNH, USNM; Hatch 1953: 158), northern Arizona (Coconino County, USNM; Snow 1906b: 162), northwestern Texas (Potter County, USNM), and Wisconsin (Purrington and Maxwell 1998: 190); also known from one specimen collected in southern Quebec (Lindroth 1969a: 1078).
This species is known from a few localities in Washington (Hatch 1953: 158) and western Oregon (Westcott et al. 2006: 8).
This species occurs from Newfoundland (Lindroth 1955a: 131) to southwestern Yukon Territory (Shpeley and Ball 1999: 422), south to the Sierra Nevada in east-central California (Papp 1978: 165; Nelson 1988a: Fig. 1), Santa Catalina Mountains in southern Arizona (Nelson 1988a: Fig. 1), and southern Colorado (Wickham 1902: 240; Lindroth 1969a: 1085); isolated on some mountains of northern New England. The records from “Vermont,” “Massachusetts,” “Connecticut,” and “Rhode Island” (Erwin et al. 1977: 4.58) are likely in error; that from Alaska (Lindroth 1969a: 1085) probably refers to
This species is known from southeastern Arizona (Bousquet and Larochelle 1993: 334) and central New Mexico (Torrance County, Ken Karns pers. comm. 2009) south to the state of Durango along the Sierra Madre Occidental (Ball and Shpeley 1992a: 61).
This Holarctic species ranges from Ireland to the Far East, south to Spain, Portugal, Greece, Kazakhstan, Mongolia, northern China (Kabak 2003: 418), and in the Nearctic Region from Alaska to northwestern Northwest Territories [see Shpeley and Ball 1999: Fig. 6].
Nineteen species in the temperate, subtropical, and tropical regions of North America (11 species) and Middle America (11 species).
Casey (1920) published a key to all but three (
Lorenz (2005: 465) listed
This species is known only from the type series.
This species is known only from the type series.
This species ranges from east-central Georgia (Emanuel County, UASM) to southern Florida (Collier County, CMNH). The record from “South Carolina” (Bousquet and Larochelle 1993: 268) needs confirmation.
This species is known only from the type locality in southeastern Arizona.
This species ranges from Maryland (Prince Georges County, UASM) and northeastern Virginia (UASM) south to central Florida (Peck and Thomas 1998: 24) and southwestern Alabama (Mobile and Washington Counties, UASM). All other state records listed in Bousquet and Larochelle (1993: 269) as well as that from southeastern Pennsylvania (Rathvon 1869: 524, as
This species ranges from Nova Scotia (Lindroth 1954c: 307) to northwestern Minnesota (Becker County, UASM), south to southeastern Texas (Brazoria County, CMNH) and central Florida (Peck and Thomas 1998: 24). The record from “North Dakota” (Bousquet and Larochelle 1993: 269) needs confirmation.
This species is known only from the type locality in central North Carolina and one locality in northern Georgia (Fattig 1949: 40).
This species ranges from southwestern Maine (Majka et al. 2011: 47) and southern Quebec (Bousquet 1987a: 133) to eastern North Dakota (Tinerella 2003: 636), south to western Texas, Nuevo León in Mexico, and southern South Carolina [see Hunting 2009: Fig. 4.12]. The record from northern Sonora (Bates 1884: 296) needs confirmation.
This species is known from western Arizona to western Texas, south to Nuevo León, Coahuila, Durango, and Sonora in Mexico (Ball and Shpeley 1992a: 62).
This species ranges from northeastern California to north-central Colorado, south to eastern Michoacán in Mexico and southern California [see Hunting 2009: Fig. 4.23].
This species is known only from the type locality in southern Arizona.
About 115 species (Lorenz 2005: 464-465) in the Nearctic (eight species of
1. The family-group names
Seventy-five species in the Western Hemisphere arrayed in two subgenera:
Sixty-seven species in the temperate, subtropical, and tropical areas of the Nearctic (nine species) and Neotropical (64 species) Regions.
Horn (1882: 156) wrote a key to four of the species found in North America. A taxonomic revision of the subgenus is needed.
This species is known from Alabama (Löding 1945: 22) and northwestern Georgia (Fattig 1949: 41) south to southern Florida (Peck and Thomas 1998: 23).
This species is known from Monroe and Dade Counties in southern Florida (Thomas 2011: 308), Cuba, Jamaica, and the Grand Cayman Island [see Ball and Shpeley 2009: Fig. 9].
This species is known from the Mexican states of Morelos, Colima (Bates 1891a: 271) and Baja California Norte (Notman 1919b: 234) and from southern Arizona (Ober and Maddison 2008: 30).
This subspecies ranges from Massachusetts (Davidson et al. 2011: 514) to southeastern Minnesota (Gandhi et al. 2005: 932), south to southern Texas (Live Oak County, Brian Raber pers. comm. 2010), northwestern Louisiana (Natchitoches Parish, USNM), and central Florida (Highlands and Pasco Counties, CMNH; Horn 1882: 157); also known from southern Arizona (Pima and Santa Cruz Counties, UASM, CMNH) and the Baja California Peninsula (Horn 1895: 226).
The subspecies
This species ranges from southern and western Texas (Duval, Val Verde, and Jeff Davis Counties, CMNH) to southeastern California (Dajoz 2007: 20) and the Baja California Peninsula (LeConte 1867b: 364).
This species is found along the Coastal Plain from South Carolina (Ciegler 2000: 120) to southern Florida (Peck and Thomas 1998: 23), west to southern Alabama (Löding 1945: 22); also recorded from the Bahamas (Turnbow and Thomas 2008: 11).
This species ranges from southern Arizona (Bousquet and Larochelle 1993: 334) to Panama (Bates 1883a: 189).
This subspecies is known from Dade County in southeastern Florida (Thomas 2011: 307), the Bahamas, Cuba, and the Cayman Islands [see Ball and Shpeley 2009: Fig. 13].
Two other subspecies,
The range of this species extends from Rhode Island (USNM) to northeastern Kansas (Popenoe 1877: 23), including southernmost Ontario (Lindroth 1969a: 1088), south to southern Texas (Wickham 1897: 112; Galveston and San Patricio Counties, UASM) and the Florida Keys (Peck and Thomas 1998: 23).
Worldwide, with about 735 species arrayed in 48 genera (Lorenz 2005: 470-478, 480, as
Jeannel (1949a: 991) grouped this subtribe and the demetriadines under the subfamily
About 105 species (Lorenz 2005: 475-477) in the Nearctic (two species), Neotropical (20 species), Australian (one adventive species in New Zealand), Oriental, Palaearctic (53 species), and Afrotropical Regions. These species are placed in three subgenera:
About 50 species in the Nearctic (two species, one of them adventive), Neotropical (20 species), Australian (one adventive species in New Zealand), Oriental (four species), and Palaearctic (28 species) Regions.
Larson (1998: 126) discussed the structural differences between the two species found in North America.
This European species is adventive in North America where it is known only from three specimens collected in Newfoundland (Larson 1998: 126) and Nova Scotia (Majka and Klimaszewski 2004: 9; Halifax County, UASM). The first inventoried specimen collected on this continent was found in 1952 in Armdale, near Halifax, Nova Scotia (UASM).
The range of this species extends from Cape Breton Island (Bousquet 1987d: 107) to Vancouver Island (Lindroth 1969a: 1043), south to southern California (Fall 1901a: 48; MCZ), the states of Sonora (CNC) and Durango (CNC) in Mexico, and the Florida Panhandle (Peck and Thomas 1998: 24). Fossil remnants, undated but believed to be younger than 3 million years B.P., have been unearthed in northwestern Yukon Territory (Elias and Matthews 2002: 914).
Lindroth (1969a: 1042-1043) studied superficially the structural variation in this species and concluded there was a single species but that the specimens west of the Rockies “may very well be treated” as a subspecies for which the name
About 50 species (Lorenz 2005: 477) in the Nearctic (two species, one of them adventive), Oriental (two species), Palaearctic (about 35 species, 15 of them endemic to the Canary and Madeira Islands), and Afrotropical (11 species) Regions.
Bousquet (2004b: 49-51) commented on the structural differences between the two species found in North America.
This Coastal Plain species is found from “Massachusetts” (Lindroth 1969a: 1043) to southern Florida (Peck and Thomas 1998: 24), west to Louisiana (LeConte 1880b: 163) and north to western Tennessee (Tipton County, CMNH).
This Palaearctic species is adventive in North America where it known from four specimens (CNC) collected in 1996 on Vancouver Island, at Saanich Peninsula, Island View Beach (Bousquet 2004b: 49).
About 130 species (Lorenz 2005: 473-474) in the Nearctic (eight species), Neotropical (one species endemic to Cuba and one species in northern Mexico), Oriental, Palaearctic (about 55 species), and Afrotropical Regions.
Lindroth (1969a: 1047-1054) reviewed the Nearctic species. Since then, one new North American species (
This subspecies is known from scattered localities from southernmost Ontario (Lindroth 1969a: 1054) to southern Arizona (Pima County, CMNH), southwestern Texas (Val Verde County, CNC), northwestern Alabama (Walker County, CNC), and southwestern North Carolina (Macon County, CMNH). One specimen simply labeled from California is also known (Lindroth 1969a: 1054).
The subspecies
This species is known from scattered localities from central Saskatchewan (Lindroth 1969a: 1053) to the Okanagan Valley in south-central British Columbia (Blades and Maier 1996: 66), south to the San Francisco area (Casey 1920: 270), northern Utah (Lindroth 1969a: 1053), northeastern Colorado (Bell 1971: 47), and southwestern South Dakota (Kirk and Balsbaugh 1975: 38).
This species is known only from the original two specimens collected at Berkeley, on the San Francisco Bay in western California.
The range of this species extends from Nova Scotia to southwestern British Columbia, though not quite reaching the coast (Lindroth 1969a: 1051), south to northern California (Lindroth 1969a: 1051; Casey 1920: 269, as
This species has passed under the name of
This subspecies is known from southern California (Fall 1901a: 48; Mateu 1974: 264), “Nevada” (Horn 1882: 134), southeastern Colorado (Bent County, CNC), and central Texas (Casey 1920: 270). The records from northeastern Kansas (Popenoe 1877: 23) and “Oregon” (Horn 1882: 134) need confirmation. The previous records from “Colorado” (Wickham 1902: 239; Mateu 1974: 264) were probably based on a misinterpretation of the type locality.
The subspecies
This species is known from a few specimens collected along the coast in southwestern British Columbia, west-central Washington (Lindroth 1969a: 1049), and Oregon (Lane County, CNC).
The range of this species extends from Vancouver Island to eastern South Dakota (Kirk and Balsbaugh 1975: 38; French et al. 2004: 557), south to southern Colorado (Wickham 1902: 239; Boulder County, CMNH), northern Arizona (Coconino County, CMNH), and southern California (Lindroth 1969a: 1053). The record from “New Mexico” (Wickham 1896c: 135) needs confirmation.
This species is found from Long Island, New York (Notman 1928: 238) to eastern South Dakota (Kirk and Balsbaugh 1975: 38), including southernmost Ontario (Lindroth 1969a: 1054), south to southeastern Texas (Horn 1882: 135; Snow 1906a: 141; Casey 1920: 270), east-central Mississippi (Snodgrass and Cross 1983: 17), and northeastern Tennessee (Hylton 1980: 25). The record from the District of Columbia (Ulke 1902: 7) needs confirmation.
About 60 species (Lorenz 2005: 472) in the Nearctic (13 species), Neotropical (five species in Middle America and the West Indies), Australian, Oriental, Palaearctic (27 species), and Afrotropical Regions.
Casey (1920: 272-276) wrote a key to all but one (
This species is known only from the type locality in central North Carolina and two localities in northern Georgia (Fattig 1949: 39).
This species is known only from the holotype collected in southwestern California.
This species is known only from the holotype collected along the coast of California.
The range of this species extends from southwestern Alberta to Vancouver Island, south to west-central California (Lindroth 1969a: 1046) and southwestern Colorado (Elias 1987: 634).
The range of this species extends from Cape Breton Island (Lindroth 1969a: 1046) to southern Saskatchewan (Ronald R. Hooper pers. comm. 2007), south to southern Arizona (Pima County, CMNH), central Texas (Casey 1920: 274, as
This species has passed under the name
This species is known from east-central Oregon (Grant County, UASM; Horn 1882: 134; Lindroth 1969a: 1044) to northern Colorado (Armin 1963: 179), including southern Idaho (Horning and Barr 1970: 25), south to central New Mexico (Bernalillo County, CMNH), southern Arizona (Pima County, CMNH; Horn 1882: 134), and the Baja California Peninsula (Horn 1894: 310).
This species is known only from the two syntypes collected along coastal California.
This species is known only from the holotype collected in northwestern Nevada.
This species ranges from Vancouver Island (Lindroth 1969a: 1047) to northwestern Montana (Flathead County, Ken Karns pers. comm. 2009), south to southwestern New Mexico (Hidalgo County, CMNH), southern Arizona (Graham, Pinal, Cochise, Pima, and Greenlee Counties, CMNH), and northern California (Shasta and Yolo Counties, CMNH).
This species is known only from the holotype collected in western California.
The range of this species extends from Cape Breton Island (Lindroth 1954c: 307, as
This species is known only from the syntypes collected in northwestern New Mexico.
This species is known only from the type series collected in southern Arizona.
About 50 species (Lorenz 2005: 470-471) in the boreal, temperate, and tropical areas of the Nearctic (one species), Australian (one widely distributed Old World species), Oriental, Palaearctic (about 35 species), and Afrotropical Regions.
The North American species is included in Lindroth’s (1969a: 1056) monograph on the Canadian and Alaskan
The range of this species extends from Newfoundland (Lindroth 1955a: 130) to east-central Alaska (Lindroth 1969a: 1056), south to the Sierra Nevada in California (Dajoz 2007: 16), southeastern Arizona (Graham and Greenlee Counties, UASM), central New Mexico (Fall and Cockerell 1907: 160), south-central South Dakota (Kirk and Balsbaugh 1975: 38), central Illinois (Wolcott 1895: 309), and western North Carolina (Yancey County, USNM) along the Appalachians. The records from “Nebraska” and “Texas” (Bousquet and Larochelle 1993: 274) need confirmation.
Fifteen species in the temperate, subtropical, and tropical areas of the Nearctic (seven species) and Neotropical (nine species) Regions.
There is no modern taxonomic revision of the species of
This species ranges from “Maine” (Larochelle and Larivière 1990a: 32) to “Washington” (Hatch 1953: 156), north to the southern part of the Prairie Provinces (Bousquet 1987a: 133), south to southwestern California (Casey 1920: 266, as
This species is known only from the holotype collected in central Washington.
This species ranges from western Idaho (Washington County, CNC) to northern Oklahoma (Alfalfa County, CMNH), south to Guatemala (Bates 1883a: 193) and southwestern California (LeConte 1851: 175; Fall 1901a: 48).
This species is known only from the type series collected in west-central California.
This species is known only from the holotype collected in northwestern Washington.
This species is known only from Utah and Washington Counties in Utah (Tanner 1928: 270).
This species is known from the type locality in west-central Washington (Hatch 1953: 156) and from southeastern Oregon (Westcott et al. 2006: 6).
Worldwide, with about 800 species arrayed in 25 genera (Lorenz 2005: 481-489). The North American fauna has 49 species (about 6 % of the world fauna) placed in two genera.
Casale’s (1998: Fig. 91) cladistic analysis suggests that this subtribe is possibly the sister-group to {cymindidines + dromiusines + physoderines + agrines + metallicines + calleidines + demetriadines + peliocypadines}.
Worldwide, with about 740 species (Lorenz 2005: 481-488) arrayed in 17 subgenera. The North American fauna includes 48 species (about 6.5 % of the world fauna) in the boreal, temperate, and subtropical regions. These species are currently placed in four subgenera.
Madge (1967) revised the North American species. Since the publication of his work, one of his species (
Twenty-five species in the Nearctic (eight species) and Neotropical (18 species) Regions.
This species ranges from southern Manitoba to eastern Oregon (Baker County, James R. LaBonte pers. comm. 1992), as far north as central Alberta, south to northern Sonora (Bates 1884: 298) and southwestern Texas [see Madge 1967: Fig. 141]. On specimen is known from Strafford County, northeastern Virginia (Hoffman 2010: 23), probably resulting from anthropogenic transport.
This species ranges from “Maine” (Larochelle and Larivière 1990a: 32) to southwestern Montana, north to northern Alberta (Fort McMurray area, Gerald J. Hilchie pers. comm. 2009), Saskatchewan and southern Manitoba, south to central New Mexico (Fall and Cockerell 1907: 159), central Texas, and northeastern Florida [see Madge 1967: Fig. 117]. The records from “California” (Erwin et al. 1977: 4.61; Bousquet and Larochelle 1993: 274) and Mexico (Chaudoir 1871a: 143) are likely in error.
This species is known from southern Arizona and western Texas (Madge 1967: 159).
This species is found east of the Rocky Mountains from southern Quebec (Larochelle 1975: 91) to northeastern North Dakota (Tinerella 2003: 637), south to southeastern Texas along the Rio Grande and southern South Carolina, west to western Texas and western Oklahoma [see Madge 1967: Fig. 137]. The records from Baja California (Horn 1894: 310), Arizona (Wickham 1898: 300; Snow 1907: 142, as
This species is known so far only from southern Arizona (Madge 1967: 159).
This species is known from southern Arizona and western Texas (Madge 1967: 158), including southwestern New Mexico (Luna County, Ken Karns pers. comm. 2009).
This species ranges from southern Arizona to western Texas (Madge 1967: 161).
This species occurs from Nova Scotia (Halifax and Colchester Counties, CNC) to eastern Minnesota, north to southern Manitoba (CNC), south to northern Oklahoma (Alfalfa County, Foster F. Purrington pers. comm. 2010), southern Louisiana, and central Florida [see Madge 1967: Fig. 126]. Old specimens simply labeled from “Texas” (Madge 1967: 157) are known.
One species in southern United States and northeastern Mexico.
This species is known from a few specimens collected in western Texas (Madge 1967: 164), southeastern Arizona (Cochise County, CNC, UASM), and “Tamaulipas” (MCZ, collection LeConte) and Nuevo León (UASM) in Mexico. The record from Baja California (Horn 1894: 310, as
Fourteen species in the Nearctic (one species) and Palaearctic (13 species) Regions.
This species ranges from the Prairie Provinces, as far north as central Alberta, south to northern Colorado (Wickham 1902: 239) and eastern Kansas (Popenoe 1877: 23), east to west-central Indiana (Downie 1957: 116; Schrock 1985: 351), west to northwestern Idaho [see Madge 1967: Fig. 120]. The record from northeastern New Mexico (Fall and Cockerell 1907: 159) needs confirmation.
Worldwide, with about 525 species (Lorenz 2005: 481-488) of which 38 (about 7 % of the world fauna) occur in North America.
This species ranges from northwestern Vermont to eastern South Dakota (Kirk and Balsbaugh 1975: 37), including southernmost Ontario (Bousquet 1987a: 133), south to Guatemala (Bates 1883a: 229, as
This species ranges from southeastern Texas (Madge 1967: 172) to Colombia (Martínez 2003: 14); also recorded from the Bahamas (Turnbow and Thomas 2008: 13), Cuba (Gundlach 1891: 16), Jamaica (Darlington 1941a: 14), Puerto Rico (Wolcott 1936: 188), and Dominica (Peck 2006: 176).
This species ranges from New Jersey to northeastern Kansas (Popenoe 1877: 23), south to Nicaragua (Bates 1883a: 240) and southern Florida, west to northwestern Texas [see Madge 1967: Fig. 119]; also recorded from Cuba (Gundlach 1891: 16; Darlington 1934: 113) and Jamaica (Darlington 1941a: 14). The record from “Wisconsin” (Bousquet and Larochelle 1993: 276) needs confirmation.
This species ranges from central Idaho and northern Oregon south to southern California along the Mexican border [see Madge 1967: Fig. 133].
The range of this species extends from northern New Jersey to north-central Colorado, north to southeastern Michigan, south at least to southern Arizona, Guanajuato and the Federal District in Mexico (Bates 1883a: 241), and central Georgia [see Madge 1967: Fig. 124]. At least one old specimen simply labeled from Wisconsin is known (Madge 1967: Fig. 124). The record from “Massachusetts” (Leng and Beutenmüller 1893: 142) needs confirmation.
This rarely collected species is known from southern Ohio (Purrington et al. 1999: 48) and southern Indiana, from North Carolina to southern Florida [see Madge 1967: Fig. 136], from southeastern Louisiana (Saint Tammany Parish, Igor M. Sokolov pers. comm. 2009), from east-central Texas (Riley 2011), and from western Arkansas (Polk and Garland Counties, Robert L. Davidson pers. comm. 2012). The records from New York (LeConte 1846b: 195), Alabama (Löding 1945: 21), and Cuba (Darlington 1934: 113) need confirmation.
This species is known from southern Arizona and the Baja California Peninsula (Madge 1967: 202).
This species ranges from southeastern Texas (Madge 1967: 212) along the Rio Grande south at least to Guatemala (Bates 1883a: 231).
This species is known from northwestern and south-central Louisiana (Caddo and Saint Landry Parishes, Igor M. Sokolov pers. comm. 2009), eastern Texas (Madge 1967: 212; Riley 2011), and “Mexico” (Erwin et al. 1977: 4.62); also recorded from the Bahamas (Turnbow and Thomas 2008: 13) and Cuba (Mateu 1977: 378). The record from South Dakota (Kirk and Balsbaugh 1975: 38) needs confirmation.
This species is found from Nova Scotia to western Washington, as far north as southern Manitoba and southwestern British Columbia (Lindroth 1969a: 1036), south to west-central California, east-central Texas (Riley 2011), and southern Florida [see Madge 1967: Fig. 127]. The species has been rarely collected in the Great Plains.
The range of this species extends from southern Alberta (Drumheller, CNC) and southern British Columbia south to southern California, southern New Mexico [see Madge 1967: Fig. 134], and western Texas (Jeff Davis County, Ken Karns pers. comm. 2009). One old specimen simply labeled from Kansas is known (Madge 1967: 201).
This species is known only from southeastern Texas (Madge 1967: 203).
This species ranges from southeastern Maine (Majka et al. 2011: 47) to eastern North Dakota (Tinerella 2003: 637), south to eastern Texas and central Florida [see Madge 1967: Fig. 135].
This species ranges from Cape Breton Island, Nova Scotia (Christopher G. Majka pers. comm. 2007) to northwestern South Dakota (Kirk and Balsbaugh 1975: 37), including northern Minnesota, south to eastern Texas and central Florida (Peck and Thomas 1998: 24) [see Madge 1967: Fig. 128].
This species is known only from a few specimens collected in southern Arizona (Madge 1967: 207).
This species is known from southern Arizona to western Texas (Madge 1967: 206), south to Guatemala (Bates 1883a: 230).
This species is known only from a few specimens collected in southeastern Texas (Madge 1967: 213).
This species is known only from Dade, Monroe, and Walton Counties in Florida (Peck and Thomas 1998: 24).
This species is found from northern New Hampshire to central Alberta, south to central California, the state of Hidalgo in Mexico (Ball and Shpeley 1992a: 64), and southern Florida [see Madge 1967: Fig. 123].
This species ranges from Nova Scotia (Lindroth 1954c: 307) and Prince Edward Island (Majka et al. 2008: 133) to southeastern Alberta, south to southeastern Texas along the Rio Grande and southern Florida. The species is also known from one locality in southwestern British Columbia and one in western Washington [see Madge 1967: Fig. 132].
This species ranges from southern Arizona and southern Texas (Madge 1967: 186) south at least to the Yucatán Peninsula (Bates 1883a: 230) in Mexico.
This species is found from Arizona (Madge 1967: 184) to western Texas (Graves and Suter 1979: 6).
The range of this species extends from southwestern Maine (Majka et al. 2011: 47) and southwestern Quebec (LeSage 1996: 22) to southeastern South Dakota (Kirk and Balsbaugh 1975: 37), including southernmost Ontario (Pettit 1869: 107; CNC), south to southeastern Texas along the Rio Grande and southern Florida [see Madge 1967: Fig. 122]. The record from “North Dakota” (Bousquet and Larochelle 1993: 278) needs confirmation.
The range of this species extends from southern Arizona to western Texas (Madge 1967: 175). The record from “Nevada” (Bousquet and Larochelle 1993: 276) needs confirmation.
This species ranges from southern Saskatchewan to south-central British Columbia, south to southern California along the Mexican border and northern New Mexico [see Madge 1967: Fig. 129]. At least one specimen is known from “Texas” (Madge 1967: 177). The record from “North Dakota” (Bousquet and Larochelle 1993: 276) needs confirmation.
The range of this species extends from New Jersey (Smith 1890: 87; Smith 1910: 210) to Arizona, north to northwestern Ohio (Hancock County, Foster F. Purrington pers. comm. 2009), southern Michigan, and northern Illinois, south to Brazil (Chaudoir 1871a: 185) and southern Florida [see Madge 1967: Fig. 125]; also found “on all the Greater Antilles” (Darlington 1953: 11, as
This species ranges from Newfoundland (Larson and Langor 1982: 594) and southern Labrador to Vancouver Island (Lindroth 1969a: 1028-1029), south at least to northern Oregon (Tillamook and Umatilla Counties, CNC), southern Wyoming (Lavigne 1977: 46; Lincoln County, CNC), eastern Nebraska (Cuming County, Foster F. Purrington pers. comm. 2009), east-central Ohio (Usis and MacLean 1998: 67), and the District of Columbia (Ulke 1902: 7). The record from east-central Missouri (Summers 1873: 134) needs confirmation.
Madge (1967: 177) listed this form as synonym of
This species occurs from southern British Columbia, including Vancouver Island, south to southern California along the Mexican border, east to southwestern Idaho [see Madge 1967: Fig. 140].
This species extends from southwestern Quebec (LeSage 1996: 22) to south-central Saskatchewan (Hooper 1977: 51), south to northeastern Kansas (Madge 1967: 174), central Missouri (Cooper County, CMNH), and northwestern North Carolina (Watauga County, CMNH). The records from the Organ Mountains in southern New Mexico (Fall and Cockerell 1907: 160) and “Mexico” (Blackwelder 1944: 55) are probably in error.
This species is known only from a few localities in southeastern Texas (Madge 1967: 173).
This species is found from southern California to western Texas, north to Mesa County in Colorado (Madge 1967: 175).
This widely distributed species ranges from Nova Scotia to northwestern Yukon Territory, south to southern California along the Mexican border, Guatemala (Chaudoir 1871a: 192), and southern Florida [see Madge 1967: Fig. 139]; also reported from some islands of the Greater Antilles (Blackwelder 1944: 56). According to Shimonoya (2004), this species has been recently found in Fukui Prefecture, Japan.
This species ranges from southern Arizona (Madge 1967: 194) to Guatemala (Bates 1883a: 240).
This species is found in southern Arizona (Madge 1967: 189). At least one specimen labeled from “Texas” is known.
Madge (1967: 189) retained this name only for the Arizona (and possibly Texas) populations but pointed out that populations, probably representing the same species but with different color patterns, were seen from Mexico and Colombia.
This species is known only from a few localities in Arizona (Madge 1967: 196).
This species is found from New Hampshire to northeastern Kansas (Knaus 1903: 188), south to eastern Texas and northern Florida [see Madge 1967: Fig. 121]. The state record of “New Mexico” (Bousquet and Larochelle 1993: 277), based on two specimens in Horn collection (MCZ) labeled “N.M.,” needs confirmation.
This species is found east of the Rocky Mountains from southern Nova Scotia to southwestern Saskatchewan, south to southeastern Texas along the Rio Grande and southern Florida [see Madge 1967: Fig. 131]; also recorded from Cuba (Darlington 1934: 113) and “Mexico” (Lindroth 1969a: 1031). The record from southwestern New Mexico (Fall and Cockerell 1907: 160, as
This species is found from Nova Scotia to southeastern British Columbia (Lindroth 1969a: 1033), south to southern California, southwestern New Mexico (Fall and Cockerell 1907: 160), southeastern Texas, and southern Florida [see Madge 1967: Fig. 118]; also recorded from “Mexico” (Bates 1883a: 240) and the state of Tabasco (Bates 1891a: 274, as
Seven species are known (Erwin 2004: 33) in the Neotropical Region, one of which extends into southern Texas.
Reichardt (1973) reviewed the species then known and published a key for their identification. Subsequently, two new species were described by Erwin (2004).
This species ranges from southern Texas to Costa Rica (Erwin 2004: 37).
Worldwide, with about 660 species arrayed in 46 genera (Lorenz 2005: 491-498). The North American fauna is represented by 24 species (about 3.5 % of the world fauna).
Erwin (2004: 6) advocated combining calleidines with agrines into one subtribe (
Seventeen species (Lorenz 2005: 495) in the Nearctic (four species), Neotropical (14 species), and Australian (one species from New Caledonia) Regions. One species,
Horn (1882: 145) provided a key for the identification of the North American species. Since then, one new species (
Fourteen species in the temperate, subtropical, and tropical areas of the Nearctic (four species, all of them extending into the West Indies or Mexico) and Neotropical Regions.
This species ranges from southeastern Georgia (Fattig 1949: 40; McIntosh County, CNC) to southern Florida (Peck and Thomas 1998: 24), west to southwestern Alabama (Leng 1915: 588; Mobile County, MCZ, USNM); also recorded from the Bahamas (Turnbow and Thomas 2008: 13). The record from Guana Island (Valentine and Ivie 2005: 275) in the British Virgin Islands needs confirmation.
This species is known only from southern Florida (Peck and Thomas 1998: 24), Cuba (Darlington 1934: 117), and Navassa (Steiner 2008: 132).
According to Ball (in Peck 2005: 39),
This species is known from northern Georgia (Fattig 1949: 40), the Florida Peninsula, as far south as Dade County (Peck and Thomas 1998: 24), and the Bahamas (Darlington 1953: 13).
The range of this species extends from southeastern New Hampshire (Strafford County, Ross T. Bell pers. comm. 2008) to southern Wisconsin (Messer 2010: 44) and northern Iowa (Wickham 1911b: 7), including southern Ontario (Pettit 1869: 107), south to central Texas (Haldeman 1852: 373; Blanco County, CMNH), southern Louisiana (East Baton Rouge, Iberville, Saint Martin, and Saint Tammany Parishes, Igor M. Sokolov pers. comm. 2009), and central Florida (Pinellas County, CMNH; Schwarz 1878: 435), west along southwestern United States to southern California (Moore 1937: 12; Kern and Fresno Counties, CNC) and the Baja California Peninsula (Horn 1894: 310).
Two Neotropical species, one of them subcosmopolitan, and one New Caledonian species (
Lindroth (1969a: 1066) stated that this species is almost cosmopolitan and probably of South American origin. In North America, it is known from “Massachusetts” (Harris 1833: 566, as
Two Nearctic species, one of them extending into northern Mexico.
Larson (1969) revised the species.
This subspecies occurs from northern California to eastern Colorado, south to southeastern Texas, Durango and Sinaloa in Mexico, and southern California [see Larson 1969: Fig. 63]. The record from “Oregon” (Bousquet and Larochelle 1993: 280) needs confirmation.
The range of this subspecies extends from southern Saskatchewan (several localities, CNC) and southern Alberta south to southwestern Idaho, northern Utah, and “Colorado” [Larson 1969: 62, Fig. 63].
This species is restricted to the Gulf Coast of Texas [see Larson 1969: Fig. 62].
About 275 species primarily in tropical areas of the Neotropical, Oriental, and Afrotropical Regions (over 95 % of the world fauna) with a few elements found in the Nearctic (11 species) and Palaearctic (ten species in Asia) Regions. The species are arrayed in three subgenera:
About 175 species confined to the Western Hemisphere. Only 11 species are endemic (five species) or extends (six species) into the Nearctic Region.
There is no key for the identification of the North American species.
This species is known from southeastern Texas (Schaeffer 1905: 142; MCZ) south at least to Veracruz, Mexico (Bates 1883a: 212); it may occur much further south, as far as Venezuela (Schaeffer 1910: 397).
This species ranges from south-central North Dakota (Burleigh County, Donald P. Schwert pers. comm. 1989) to southeastern Virginia (Hoffman and Roble 2000: 40), south to southern Florida (Peck and Thomas 1998: 24), the Bahamas (Darlington 1953: 11), and Nicaragua (CMNH), west to southern Arizona (Santa Cruz and Pima Counties, CMNH, UASM).
This species is found in southern Texas (Bousquet and Larochelle 1993: 334) south at least to Veracruz (Bates 1883a: 212) in Mexico.
This species ranges from southern South Carolina (Ciegler 2000: 126) to southern Florida (Peck and Thomas 1998: 24), west to “Texas” (Leng 1915: 587) and central Oklahoma (Hatch and Ortenburger 1930: 11). The records from Colorado (Wickham 1902: 240) and northeastern Kansas (Knaus 1901: 110) need confirmation.
This species is known only from the holotype collected in central Florida.
This species ranges from the Rio Grande Valley in southern Texas (Wickham 1897: 111) south to Guatemala and Belize (Bates 1883a: 208, as
This species ranges from southwestern California (Moore 1937: 12) to southeastern Texas (Cameron County, CNC), including northern Sonora (Bates 1884: 298); also known from Baja California Sur (Santa Rosa, MCZ).
The range of this species extends from Maine (Dearborn and Donahue 1993: 8; Foss 2001: 14) to southern Manitoba (Bousquet 1987a: 133), south to “Kansas” (Horn 1882: 141) and northwestern South Carolina (Anderson County, Robert L. Davidson pers. comm. 2012). The records from “Texas,” “Florida” (Erwin et al. 1977: 4.59), “Louisiana” (Chaudoir 1873b: 153), “Arkansas” (Bousquet and Larochelle 1993: 281), northern Alabama (Löding 1945: 22), and southwestern Georgia (Fattig 1949: 40) need confirmation.
This species is known from the Baja California Peninsula (Horn 1894: 361, as
This species ranges from “New Hampshire” (Lindroth 1969a: 1062) to northeastern South Dakota (Kirk and Balsbaugh 1975: 38), north to southern Manitoba (Lindroth 1969a: 1062; Bousquet 1987a: 133), south to eastern Kansas (Popenoe 1877: 23), central Louisiana (Rapides Parish, Igor M. Sokolov pers. comm. 2009), and northern Florida (Peck and Thomas 1998: 24). The records from Colorado (LeConte 1846b: 188; Snow 1877: 17) need confirmation.
This species ranges from southwestern New York (Notman 1928: 239) to eastern New Mexico (Chaves and De Baca Counties, CMNH), including southwestern Iowa (Fremont County, Doug A. Veal pers. comm. 2009), south to southeastern Texas (Wickham 1897: 111), southern Louisiana (East Baton Rouge, Iberville, Pointe Coupee, Saint Martin, and Saint Tammany Parishes, Igor M. Sokolov pers. comm. 2009), and southern Florida (Peck and Thomas 1998: 24). The records from “Utah,” “Arizona,” and “California” (Csiki 1932b: 1449) are likely in error.
Four species comprised this genus: one (
Larson (1969) revised the species.
This taxon is ranked as a subgenus of
This species ranges from Arizona south to the Isthmus of Tehuantepec [see Larson 1969: Fig. 58].
This species ranges from eastern Colorado and Kansas south to northeastern Mexico and southwestern Texas [see Larson 1969: Fig. 58]. The record from South Dakota (Kirk and Balsbaugh 1975: 38) needs confirmation. One specimen labeled from “Alabama” and two specimens from California seen by Larson (1969: 32) are probably mislabeled.
The range of this subspecies extends from southern Manitoba to south-central British Columbia, south to northern Oregon, Colorado (Larson 1969: 42-43), and east-central Kansas (Knaus 1907: 233). The records from “New Mexico” and “Utah” (Bousquet and Larochelle 1993: 281), based on misplaced dots on Larson’s (1969) figure 59, are in error; those from “Oklahoma” (Arnold 2008) and “Texas” (Casey 1913: 174, as
This subspecies ranges from northern Washington to eastern Wyoming (Lavigne 1977: 45, as
This subspecies is known only from south-central Oregon and northern California [see Larson 1969: Fig. 59].
This subspecies is known only from the San Francisco Bay area in California [see Larson 1969: Fig. 59].
One Nearctic species extending into northern Mexico.
The character states of the sole species are described in detail by Larson (1969: 43-44).
This species is known from “Nevada” and the Lake Tahoe region in the Sierra Nevada south to the Mexican border in California [see Larson 1969: Fig. 61]. According to Ball and Bousquet (2000: 114), the species is also found in northwestern Mexico.
Ten species in Central America and South America, of which two reach southern United States.
This genus has not been revised in modern times and such work is needed.
This species ranges from southeastern Texas (Cameron County, Edward G. Riley pers. comm. 2009) south to the Amazonia (Chaudoir 1873b: 166). This is a
This species is known only from Brevard County in central Florida (Peck and Thomas 1998: 24).
Seven Neotropical species, of which one reaches southeastern Texas.
This genus has not been revised in modern times and such work would be useful.
This species is found from southeastern Texas (Wickham 1897: 108; Snow 1906a: 141 as “
About 585 species (Lorenz 2005: 498-502) belonging to the Western Hemisphere genus
About 585 species in the Neotropical Region, one of them extending into southeastern Texas.
1. Erwin (1982a: 45) postulated that
This subspecies ranges from southeastern Texas (Wickham 1897: 106) to Guatemala (Bates 1883a: 247). As mentioned previously, the Texas specimens actually belong to an undescribed species.
The subspecies
About 70 species (Lorenz 2005: 490) in the Nearctic (three species), Neotropical (16 species of
This subtribe came out as the sister-group to {
Sixteen species restricted to the Western Hemisphere, currently arrayed in two subgenera:
Shpeley (1986) revised the species and provided a key for their identification.
Fifteen species in the Neotropical Region, of which three extend into southern United States.
This species is found from southern California and southeastern Texas south to Guatemala and southern Baja California [see Shpeley 1986: map 4].
This species ranges from southeastern Arizona south to southern Mexico [see Shpeley 1986: map 3].
This species is found along the east coast of Florida, including the Keys, and in Cuba [see Shpeley 1986: map 6].
One genus endemic to the Western Hemisphere.
Jeannel (1949a: 860) and Basilewsky (1984: 527) considered this taxon as a masoreimorph (i.e., masoreines and cyclosomines). Ball and Bousquet (2000: 113-114) retained it as a lebiines, as previously done by Ball (1960b: 158) and Lindroth (1969a: 1014), but recognized that its position remains to be determined.
Nine species in the temperate and tropical areas of the Nearctic (two species) and Neotropical (eight species) Regions.
This genus has not been revised in modern times and such work is needed.
The range of this species extends from the District of Columbia (CMNH) to “Illinois” (LeConte 1853c: 378), including southwestern Pennsylvania (Westmoreland County, Robert L. Davidson pers. comm. 2008), south to southern Texas (Cameron, Colorado, and Sabine Counties, USNM, Brian Raber pers. comm. 2010) and northern Florida (Peck and Thomas 1998: 24). The record from “Iowa” (Bousquet and Larochelle 1993: 279) needs confirmation.
This species ranges from southern Texas (Cameron and Hidalgo Counties, Edward G. Riley pers. comm. 2009; Brian Raber pers. comm. 2010) south to Guatemala (Bates 1883a: 173). This is a
Worldwide, with about 290 species (Lorenz 2005: 504-507, excluding
The planetines are included in this tribe by Lorenz (2005: 507). However Basilewsky (1963b), Reichardt (1967), Ball (1985), and Baehr (1986) presented arguments suggesting that planetines are probably more closely related to galeritines than to zuphiines.
Worldwide, with about 200 species placed in 14 genera. The North American fauna is represented by 18 species (about 9% of the world fauna).
Worldwide, with about 75 species of which six are found in the temperate and subtropical areas of North America.
Mateu (1981) reviewed the North American species and provided a key for their identification.
This species ranges from New Jersey (Morris County, USNM) to southern South Dakota (Kirk and Balsbaugh 1975: 39), including southernmost Ontario (Lindroth 1969a: 1090), south to southeastern Texas (Snow 1906a: 141; Cameron, Kleberg, and Gonzales Counties, CMNH, MCZ) and the Florida Keys (Peck and Thomas 1998: 25), west to southern Arizona (Mateu 1981: 118); seemingly isolated in western Oregon (Malkin 1943: 52).
This species is known only from the original two specimens.
This species is known from western Arkansas (Polk and Hempstead Counties, CMNH, MCZ), Texas, as far south as the Rio Grande in the southeast (Mateu 1981: 125; Dajoz 2007: 18-19), southern Arizona (Pima County, MCZ), and central California (LeConte 1879c: 62). The record from “Oklahoma” (Arnold 2008) needs confirmation.
This species is known from “Louisiana” (USNM) and from Cameron (Schaeffer 1910: 396) and Zapata Counties (CMNH) in southern Texas.
This species ranges from southwestern Arizona to southeastern Texas (Mateu 1981: 127). It was also reported from “Mexico” (Chaudoir 1863: 314; Liebke 1933: 471).
This species is known from a few scattered localities from southern California to southeastern Texas (Mateu 1981: 119).
About 55 species in temperate, subtropical, and tropical areas of the Nearctic (12 species but only four endemic), Neotropical (about 40 species), and Australian (nine species in Australia) Regions.
Eighteen species (Lorenz 2005: 505, as
Two of the North American species are included in Liebke’s (1934: 372-373) key to the species of this genus. A taxonomic revision is needed.
This Coastal Plain species ranges from southeastern Virginia (Davidson 1995: 18) to central Florida (Peck and Thomas 1998: 25), west to southeastern Texas (Cameron County, USNM).
This species is known only from the type locality in southwestern California.
The range of this species extends from southern Washington (Zack et al. 2003) to southern California (Fall 1901a: 47), east along the southwest to western Arkansas (Polk County, Robert L. Davidson pers. comm. 2008) and southern Texas (Zapata County, CMNH; Liebke 1934: 373), north to central Kansas (Knaus 1905a: 218), south at least to Oaxaca (Bates 1883a: 166).
Thirty-four species (Lorenz 2005: 505) in the Nearctic (nine species but only two endemic), Neotropical (23 species), and Australian (nine species in Australia) Regions.
Messer (2011: 422-423) published a key to all species found in the United States and Mexico.
This taxon is considered as a distinct genus by some authors (e.g., Ball and Bousquet 2000: 115) but Baehr (1985: 36) found out that the character states between
This species is found in central and southern Florida (Darlington 1935a: 161; Peck and Thomas 1998: 25), the Bahamas (Turnbow and Thomas 2008: 15), Cuba (Chaudoir 1877: 252; Peck 2005: 39), and the Dominican Republic (CMNH).
This species is known from Hidalgo and Cameron Counties in southern Texas (Peter W. Messer pers. comm. 2011), from southern Florida (Bousquet and Larochelle 1993: 334), and from Cuba (Darlington 1934: 128), the Cayman Islands, and Dominican Republic (Messer 2011: 422).
The range of this species extends from the District of Columbia (Ulke 1902: 7) to western South Dakota (Kirk and Balsbaugh 1975: 39), south to southern Texas (Johnson 1978: 68) and southern Florida (Peck and Thomas 1998: 25); also recorded from the Bahamas (Turnbow and Thomas 2008: 15), Cuba (Darlington 1934: 128), Jamaica (Darlington 1941a: 14), Guana Island in the British Virgin Islands (Valentine and Ivie 2005: 275), Dominican Republic, and Mexico (Messer 2011: 423). The record from Arizona (Snow 1906b: 162) is probably in error.
This species is known from Horn Island in southeastern Mississippi (Drew A. Hildebrandt pers. comm. 2010), “Texas” (Liebke 1934: 374), and Veracruz in Mexico (Bates 1883a: 166).
This species ranges from southeastern California (Fall 1901a: 47; Andrews et al. 1979: 28) and west-central Nevada (Storey County, David H. Kavanaugh pers. comm. 2008) east to southeastern Texas (Cameron County, CMNH, MCZ; LeConte 1880a: 85; Snow 1906a: 141), south to the states of Veracruz and Nayarit in Mexico (Peter W. Messer pers. comm. 2011); also recorded from “Oklahoma” (Messer 2011: 423).
This species is known only from the type locality in southwestern California.
This species is known from Veracruz in Mexico and from two specimens collected in Live Oak County in southern Texas (Messer 2011: 420).
This species ranges from southeastern Virginia (Davidson 1995: 18) to eastern Missouri (Anonymous 2007), south to southern Texas (Johnson 1978: 68) and central Florida (Peck and Thomas 1998: 25); also recorded from Cuba (Liebke 1934: 373; Peck 2005: 39).
This species ranges from southeastern Oregon (LaBonte 1996: 357) to the Baja California Peninsula (Horn 1897: 367) and southern Arizona (Maricopa and Pima Counties, MCZ); its is also known from western Texas (El Paso and Brewster Counties, Robert L. Davidson pers. comm. 2012).
Worldwide (though not represented in Europe), with about 130 species (Lorenz 2005: 507-509, as
About 105 species in the temperate, subtropical, and tropical areas of the Nearctic (eight species of
Eighty-two species in the Western Hemisphere (58 species) and the Old World (24 species) currently arrayed in two subgenera, both represented in the Nearctic Region. Only eight species are found in North America and a single one (
Reichardt (1967) revised the Western Hemisphere species and provided keys for their identification.
Seven species in North America (seven species), Middle America (six species), and the West Indies (one species).
Subsequent to Reichardt’s (1967) revision, Ball and Nimmo (1983) published a synopsis of the species of
This species is known from southeastern Louisiana (Saint Tammany Parish, Igor M. Sokolov pers. comm. 2009), Missouri, southeastern Nebraska (Nemaha County, Foster F. Purrington pers. comm. 2011), eastern Kansas (Popenoe 1877: 23; Horn 1872c: 385), eastern Texas, several mountain ranges in southern Arizona, and from the Sierra Huachinera of eastern Sonora and western Chihuahua in northern Mexico [see Ball and Nimmo 1983: Fig. 10]; it is also listed from “Oklahoma” by Arnold (2008). The record from eastern Iowa (Wickham 1911b: 7) needs confirmation.
The range of this eastern species extends from Rhode Island (Sikes and Webster 2005: 317) to southeastern South Dakota (Kirk and Balsbaugh 1975: 39), south to southeastern Texas and southern Florida [see Ball and Nimmo 1983: Fig. 14].
This species ranges from southern Arizona southwards along the Pacific Coast to Guerrero [see Ball and Nimmo 1983: Fig. 7].
This species is found from southern Quebec to southeastern South Dakota, north to southern Manitoba (Bousquet 1987a: 134), south to east-central Texas and central Florida; isolated populations are known from southwestern New Mexico (Grant County, Robert L. Davidson pers. comm. 2008), southeastern Arizona, and Chihuahua in Mexico [see Ball and Nimmo 1983: Fig. 10].
The range of this subspecies is disjunct: there is an eastern component ranging from Virginia to southwestern Kentucky (Trigg County, Foster F. Purrington pers. comm. 2011) southwards to southern Florida and South
The subspecies
This species ranges from southeastern Nebraska (Lancaster County, Foster F. Purrington pers. comm. 2011) to south-central Arizona, south to Costa Rica [see Ball and Nimmo 1983: Fig. 12].
This species is known from a few localities from eastern Texas southwards along the Gulf and Caribbean Coasts to Honduras [see Ball and Nimmo 1983: Fig. 7].
Seventy-five species in the Western Hemisphere (51 species) and the Old World (24 species). One Neotropical species extends into southern Texas.
This species ranges from the Rio Grande in southeastern Texas south to Colombia (Martínez 2003: 7) [see Reichardt 1967: Fig. 76]. The record from “Arkansas” (Bousquet and Larochelle 1993: 285) is in error.
Worldwide (though absent from Europe) with about 180 species (Lorenz 2005: 510-512) arrayed in 26 genera: five of these genera (about 60 species) are represented in the Western Hemisphere and 21 (about 120 species) in the Eastern Hemisphere. The group is underrepresented in the Northern Hemisphere with 14 species (about 8% of the world fauna), only five of them being endemic (about 3%). Two subtribes are recognized:
About 140 species in the Nearctic (seven species of
Twenty-three species in temperate, subtropical, and tropical areas of the Nearctic (seven species) and Neotropical (20 species) Regions.
Ball (1956a) revised the North American species. The taxonomy and nomenclature of these species have not changed since.
This species is found along the Coastal Plain and Piedmont from southern New Jersey to southern Florida (Peck and Thomas 1998: 25), west to southeastern Mississippi (Ball 1956a: 85, Fig. 34). The record from south-central Kansas (Knaus 1901: 110) is probably in error.
This species ranges from New Jersey (Smith 1890: 90; Smith 1910: 212) to northwestern South Carolina (Ciegler 2000: 130), west to the Baboquivari Mountains in southern Arizona, north to southwestern Utah (Washington County, CMNH) and northeastern Kansas, south to Sinaloa in Mexico (Ball 1956a: 78). The records from New York City and vicinity (Leng and Beutenmüller 1893: 143) and “Florida” (Leng 1920: 67) need confirmation.
This species is found from western Texas to south-central Arizona, south at least to Durango in Mexico (Ball 1956a: 84, Fig. 34). The record from “Oklahoma” (Arnold 2008) needs confirmation.
This species is found from Massachusetts to west-central Georgia (Fattig 1949: 41), west to eastern Texas (Ball 1956a: 75), and north to central Illinois (Purrington et al. 2002: 200) and western Michigan (Newaygo County, CMNH).
This species ranges from the mountains of southern Arizona to southwestern Texas, south at least to Durango (Ball 1956a: 86, Fig. 34).
This subspecies occurs from southeastern New Hampshire (Cooper 1976: 165) to southeastern Wyoming (Platte County, CMNH) and northern Colorado (Ball 1956a: Fig. 33), south to Oklahoma (French et al. 2001: 228; Grady County, CMNH), west-central Arkansas (Garland County, Robert L. Davidson pers. comm. 2012), and northeastern Georgia (Fattig 1949: 42). The record from southwestern Alabama (Löding 1945: 23) needs confirmation.
This subspecies is endemic to the Florida Peninsula (Peck and Thomas 1998: 25).
This subspecies is known for sure only from Mississippi (Harrison and Lafayette Counties, Drew A. Hildebrandt pers. comm. 2008), “New Jersey,” and “Alabama” (Ball 1956a: 79) where Löding (1945: 23) recorded it from Colbert and Mobile Counties. The record from southwestern Ohio (Blatchley 1910: 155) is probably in error; those from Sand Point and Tampa in Florida (Leng 1915: 589) need confirmation.
This species ranges from southwestern Kentucky (Trigg County, Foster F. Purrington pers. comm. 2010) and central Tennessee to southeastern New Mexico (Lea and Chaves Counties, CMNH), north to southwestern Iowa (Fremont County, Doug A. Veal pers. comm. 2009) and southeastern Nebraska (Jefferson County, Foster F. Purrington pers. comm. 2010), south to northeastern Mexico (Ball 1956a: 82-83, Fig. 34) and southwestern Georgia (Fattig 1949: 42). The records from southern Arizona (Snow 1907: 142) and Indiana (Blatchley 1910: 156) need confirmation.
About 325 species (Lorenz 2005: 388-390) in the Western Hemisphere (35 species) and the Australian (285 species, close to 90% of the world fauna), Oriental (four species of
Thirty species in temperate, subtropical, and tropical areas of the Western Hemisphere (30 species) and southern Australia (three species) arrayed in two subgenera:
According to Baehr (1994: Fig. 1),
Twenty-seven species in the Western Hemisphere.
Notman (1925) published a key to all species known at the time except
This species is known only from the type locality in northern Arizona.
This species is known only from the type locality in southern New Mexico.
This species is known from central and southern Arizona (Griffith 1900: 566; Notman 1925: 22) and southwestern New Mexico (Fall and Cockerell 1907: 162; Notman 1925: 22).
This species has been cited from Butte County in southeastern Idaho (Stafford et al. 1986: 289), Churchill County in western Nevada (Bechtel et al. 1983: 474), Contra Costa County (Notman 1925: 21) and the San Joaquin Valley (Fall 1901a: 51) in California, and the state of Durango (Bates 1883a: 255) in Mexico.
This species is known from southeastern Oregon (Hatch 1953: 187), Sacramento County in California, and northwestern Utah (Notman 1925: 22).
This species is known from southeastern Arizona (Notman 1925: 20). The record from “California” (Csiki 1933a: 1638) needs confirmation.
This species is known from San Diego County in southwestern California and Yavapai County in central Arizona (Notman 1925: 19).
This species is known from Inyo (Horn 1867b: 151) and Tulare (Notman 1925: 21) Counties in California.
This species is known only from the type locality in westernmost Texas.
This Coastal Plain species is found from northern South Carolina (Ciegler 2000: 130) to Louisiana (Notman 1925: 22; Erwin and Geraci 2008: 85).
This species is known only from the holotype collected in southwestern California.
This species is known from Pima, Pinal (Notman 1925: 15), and Cochise (MCZ) Counties in southern Arizona.
This species is known from southern California (Van Dyke 1943: 30; Andrews et al. 1979: 28).
This species is known only from the holotype collected in southern Arizona.
This species is known only from the holotype collected in southern Arizona.
This species is known only from the holotype collected in southern Arizona.
This species has been reported from San Diego County in southwestern California and Lyon County in west-central Nevada (Notman 1925: 18).
This species is known only from the holotype collected in northwestern Utah.
Several persons have contributed to this catalogue. Thomas C. Barr, Jr., Ross T. and Joyce Bell, James C. Bergdahl, Eric van den Berghe, Sydney G. Cannings, Donald S. Chandler, Robert L. Davidson, Garry A. Dunn, Kamal J.K. Gandhi, Vince Golia, Gerald J. Hilchie, Drew A. Hildebrandt, Richard L. Hoffman, Ronald R. Hooper, Ken Karns, David H. Kavanaugh, William L. Krinsky, James R. LaBonte, Paul K. Lago, David W. Langor, Serge Laplante, André Larochelle, Kirk J. Larsen, Harry J. Lee, Jr., David R. Maddison, Christopher G. Majka, Peter W. Messer, Steve Nanz, Robert E. Nelson, Darren A. Pollock, Foster F. Purrington, Brian Raber, Edward G. Riley, Donald P. Schwert, Derek S. Sikes, Igor M. Sokolov, Doug A. Veal, and Reginald P. Webster provided many new state or province records. George E. Ball, Robert L. Davidson, Serge Laplante, David R. Maddison, and Gerald R. Noonan granted permission to include unpublished nomenclatural findings (new synonymies) from their research. Philip D. Perkins and Hans Silfverberg provided information on T.W. Harris’ and Mannerheim’s collections respectively, Robert E. Acciavatti, George E. Ball, Shawn M. Clark, Lee H. Herman, Fritz Hieke, Stuart Hine, and Bernd Jaeger information on type specimens, and Shun-Ichi Uéno information about K. Shirahata. Nadine Dupérré provided label data of a holotype housed at AMNH and information from books and journals available at the AMNH. Paul Skelley and Boris Kataev sent photocopies of pages of books housed in the Division of Plant Industry Library (FSCA) and the Zoological Museum of Moscow University Library respectively. Warren E. Steiner, Jr. and Terry L. Erwin, Robert L. Davidson, and Philip D. Perkins assisted by arranging visits I made in 2008 to collections at the USNM, CMNH, and MCZ respectively. George E. Ball reviewed the manuscript section on the genus
I wish to thank also Steve Gamman and Patricia Madaire of the Canadian Agricultural Library (Entomology) in Ottawa, Eileen Mathias of The Ewell Sale Stewart Library at The Academy of Natural Sciences in Philadelphia, Dana Fischer of the Ernst Mayr Library at the Museum of Comparative Zoology in Cambridge, Massachusetts, and Marty Schlabach at the Comstock Memorial Library of Entomology at Cornell University in Ithaca, New York, for their assistance in locating obscure and difficult-to-find works. I am indebted to my colleague Aleš Smetana for access to his impressive personal library.
Finally, I acknowledge Henri Goulet for the permission to use some of his nice color illustrations.
Dates of publication are important for taxonomic purposes and searches have been made to find precise dates for all works cited in this catalogue. The dates are listed after the references, unless only the year was found. They were obtained either from the book or the journal itself (in such cases no reference is given), from library stamps, bibliographic works (abbreviated references given unless the work is listed in the “reference” section), or from the following recording journals and society journals:
Dates of receipt (stamps) from several libraries were used to establish dates of publication of works in the absence of more precise dates. The year of publication was not changed if the stamp was dated in the month of January of the following year.
Months of issues for journal published at regular intervals (such as
In reference to the dates given, the following symbols and abbreviations are used:
>: after
<: before
CAL: Canadian Agriculture Library at Eastern Cereal and Oilseed Research Center, Ottawa, Ontario
CISTI: Canada Institute for Scientific and Technical Information, Ottawa, Ontario
CML: Canadian Museum of Natural History Library, Gatineau, Quebec
CUL: Cornell University, Ithaca, New York
DP: Date of publication
McD: McDonald College, Sainte-Anne-de-Bellevue, Quebec
NRC: National Research Council, Ottawa, Ontario
OTU: University of Ottawa Library, Ottawa, Ontario
USNM: Smithsonian Institution Library, Washington, D.C.
Along with the reference, the following information is provided: pages where the character states of the species are indicated, the figures illustrating character states, the stage described, and the provenance of the material, either reared (RR) or field collected (FC). A question mark in front of a reference indicates that the identification of the taxon is uncertain according to the author(s). A dagger after the name of a species indicates that the taxon is adventive in North America. The abbreviations used are:
Genus A sp. A (Florida): Garner (1954: 270, 271; Pl. 19, Figs a-c) [L2-3; FC].
A number of available species were listed originally as from North America and were subsequently found not to belong to the region. The following is an annotated alphabetical list of species described from specimens mislabeled or incorrectly assumed to be from North America.
Many species- or genus-group names have been reported from North America but are unavailable (
These asterisks were dropped in the publication of Say’s entomological works by LeConte.
These names have been credited to Eschscholtz by almost all authors I have seen although there are proper citations of Fischer von Waldheim’s plates in Eschscholtz’s work. This suggests that the plates were available before the publication of Eschscholtz’s work. However because Fischer von Waldheim’s validation of the names is through illustrations, it is always possible that Fischer von Waldheim simply gave Eschscholtz the position of these species on his forthcoming plates.
By 1825, Major LeConte had sent Dejean more than 600 species of beetles (Dejean 1825: xxv). He also visited Paris in 1828 and gave Dejean a huge [“une immense”] collection of insects from the United States (Dejean 1828: vi). All species named “
A copy of the letter was published in
Following Maddison et al. (1999: 104), the expression ‘basal grade carabids’ is restricted to lineages branching off early along the evolutionary path of the family, ‘middle grade carabids’ to the lineages placed by Jeannel (1941b) in his ‘Stylifera’ and ‘higher carabids’ to the numerous lineages currently included in the subfamily
† = adventive species in North America; ‡ = Holarctic species. The species list is alphabetic within the subgenera and does not necessary follow the catalogue listing.
All vernacular names of tiger beetles are taken from Pearson et al. (2006) and Erwin and Pearson (2008).
Several new synonyms are proposed in this subgenus. All are based on a manuscript written by Serge Laplante in the 1990s originating from a study of the type specimens of all species then considered valid and the specimens in the CNC. The new synonymies should be credited to him.
This subgenus is herein removed from synonymy with
The field-collected larva described as
Gaumer (1977) described larvae of five subspecies of
The field-collected and reared larvae described by Garner (1954: 220, 221; Pl. 14, Figs a-c) as
As pointed out by Jeannel (1920a: 510), the field-collected larva described by Xambeu (1901: 58, 59) as
The field-collected larvae described by Garner (1954: 211, 212; Pl. 13, Figs a-c) as