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Catalogue of Geadephaga (Coleoptera, Adephaga) of America, north of Mexico

PMCID: PMC3577090

PMID: 23431087

Abstract

Abstract All scientific names of Trachypachidae , Rhysodidae , and Carabidae (including cicindelines) recorded from America north of Mexico are catalogued. Available species-group names are listed in their original combinations with the author(s), year of publication, page citation, type locality, location of the name-bearing type, and etymology for many patronymic names. In addition, the reference in which a given species-group name is first synonymized is recorded for invalid taxa. Genus-group names are listed with the author(s), year of publication, page citation, type species with way of fixation, and etymology for most. The reference in which a given genus-group name is first synonymized is recorded for many invalid taxa. Family-group names are listed with the author(s), year of publication, page citation, and type genus. The geographical distribution of all species-group taxa is briefly summarized and their state and province records are indicated. One new genus-group taxon, Randallius new subgenus (type species: Chlaenius purpuricollis Randall, 1838), one new replacement name, Pterostichus amadeus new name for Pterostichus vexatus Bousquet, 1985, and three changes in precedence, Ellipsoptera rubicunda (Harris, 1911) for Ellipsoptera marutha (Dow, 1911), Badister micans LeConte, 1844 for Badister ocularis Casey, 1920, and Agonum deplanatum Ménétriés, 1843 for Agonum fallianum (Leng, 1919), are proposed. Five new genus-group synonymies and 65 new species-group synonymies, one new species-group status, and 12 new combinations (see Appendix 5) are established. The work also includes a discussion of the notable private North American carabid collections, a synopsis of all extant world geadephagan tribes and subfamilies, a brief faunistic assessment of the fauna, a list of valid species-group taxa, a list of North American fossil Geadephaga (Appendix 1), a list of North American Geadephaga larvae described or illustrated (Appendix 2), a list of Geadephaga species described from specimens mislabeled as from North America (Appendix 3), a list of unavailable Geadephaga names listed from North America (Appendix 4), a list of nomenclatural acts included in this catalogue (Appendix 5), a complete bibliography with indication of the dates of publication in addition to the year, and indices of personal names, supraspecific names, and species-group names.

Full Text

A comparison of the number of valid species and genera between this and previous checklists / catalogues is presented in Table 1.
The distributional component consists of a list of state and province records, using the same two-letter postal service style abbreviations used in the 1993 catalogue (Table 2), and a short description of the distribution, usually referring to the northeasternmost, northwesternmost, southwesternmost, and southeasternmost states or provinces. In addition, records for Cape Breton Island, the Queen Charlotte Islands, Vancouver Island, and the Channel Islands are indicated in parentheses after their respective provinces or states. Western Hemisphere countries are listed for species found south of the area covered. States and provinces placed in quotation marks in the descriptive section indicate that only the state or province was given without further precision in the reference cited. The starting point for the distributional records used in this work is Bousquet and Larochelle’s (1993) catalogue. However, many of their records were undocumented or came from old lists and were not always reliable. State and province records undocumented or considered doubtful are shown in square brackets following the accepted records. Except for the records which come from identifications generally made by Fritz Hieke, almost all records from CMNH specimens are based on identifications made by Robert L. Davidson, those from LSAM specimens on identifications made by Igor Sokolov, and those from CNC, MCZ, and USNM specimens from identifications or confirmations made by myself. The records provided by Ken Karns and Brian Raber are based on identifications made by Robert L. Davidson.
Besides those used for provinces and states (see Table 2), the following abbreviations are used in the text:
New taxa. In the xviii and first half of the xix Century it was common practice for authors not to indicate the attribution of the new species-group taxa. Instead, some authors added the word mihi after the specific name, usually to indicate a taxon that the author, himself, was describing. Several collectors provided names for their specimens, even for undescribed ones, and these specimens often circulated among European coleopterists through exchange, gift, or sale. Many undescribed species were subsequently described or illustrated under the collector’s names by different authors. For these, citations are provided in this catalogue only to the first description or illustration of each species unless the term “new species” or an equivalent expression (such as an asterisk preceding the specific epithet as in Say 1823a1) was included with the species-group name subsequently described or illustrated. Sometimes a species was described / illustrated by different authors the same year under the same names. One example concerned several species (i.e., , , , , and )2 described by Eschscholtz in 1823 in the Mémoires de la Société Impériale des Naturalistes de Moscou (volume 6) and illustrated by Fischer von Waldheim on plates available the same year (Sherborn 1922: liii), but included in his Entomographie de la Russie (volume 2) issued in 1824. In such cases, citations are given for the oldest description / illustration (for exceptions see previous entry, Principle of priority) but references to subsequent descriptions / illustrations are noted after the entry of the valid name.
LeConte is without doubt the most outstanding North American coleopterist of the xix Century, not only because he described 514 new genus-group and about 4,730 new species-group taxa of beetles (Henshaw 1882: 270), but because he was the first to work seriously on the classification of the North American fauna. During his scientific activity, which lasted almost 40 years, he described 724 new species-group taxa of Geadephaga from North America, 439 (61%) of which were not previously described. LeConte built his collection through his own collecting but also from gifts he received and identifications he provided to many persons from whom he usually retained all or some of the specimens. There is also little doubt that his father, Major John Eatton LeConte3, left his collection to his son. Evidence supporting this can be found in PageBreakLeConte (1856a: 49) when he indicated that his second specimen of “came from the old collection of my father.” LeConte was a generous man and often offered some of his specimens to visitors (such as Motschulsky [q.v.]) or sent some to acquaintances (such as Chaudoir [q.v.] and Putzeys [q.v.]) though it seems that he retained at least one specimen of each species. Unfortunately in the xix Century the type concept for species-group taxa was not developed and LeConte sometimes gave the only syntype he had in his collection and retained one or more specimens that he acquired after the original descriptions. Therefore, syntypes of some of LeConte’s species are not in his collection. Moreover, syntypes of some of his species are difficult to find in his collection. LeConte had the habit of mixing the specimens of the new species he considered later as synonyms with those of the valid species. Since many of his specimens only bear a colored disc for label, syntypes of several of his species are not readily ascertainable.
In April 1875, LeConte wrote to his friend Alexander Agassiz, director of the Museum of Comparative Zoology in Cambridge, and expressed the wish that his collection be deposited at the museum after his death4. His collection was packed and transported by his longtime friend George Horn. It now stands separate from the general collection along with that of Horn.
The current classification of the Carabidae is based mainly on morphological data of adults although molecular sequence data have been used recently to discuss various aspects of carabid phylogeny. Despite several attempts there is no consensus on the classification of several subfamilies or tribes. This is particularly evident among ‘basal grade’ carabids.5
The world classification of family-group taxa, which has been adopted for the North American fauna in this catalogue, is outlined in Table 3.
Table 4 lists the adventive species found on this continent. All but two are accidental introductions (i.e., immigrants). Several carabids were intentionally introduced in New England during the first half of the xx Century for gypsy moth ( Linnaeus) control, including Kirby, Linnaeus, Fabricius, Herbst, Linnaeus, Paykull, and Linnaeus (see Smith 1959), but only and have become established.
Most of the remaining 57 adventive species found on this continent prior to the xx Century have probably been introduced with bulk rock, bricks, mortar and no doubt soil taken aboard sailing vessels as ballast and dump ashore on this continent (Brown 1940a; Lindroth 1957c). In the xx Century, species have been introduced into North America likely through shipments of nursery stock, within the soil bagged around the roots, as suggested by Kavanaugh and Erwin (1985), Spence and Spence (1988: 166), and Bousquet (1992a). Despite that quarantine regulations were established in the 1960s for the importation of nursery stock, additional species still continue to land and become established on this continent (Fig. 1). The most recent one is in the Willamette Valley in northwestern Oregon (Kavanaugh and LaBonte 2008).
Figure 1 illustrates the arrival of adventive species in North America through time based on the collecting date of the first inventoried specimen. The rate of arrival has been steady in the xx Century.
Figure 2 illustrates through decades the number of North American species-group taxa described and the number of those regarded as valid today based on the present catalogue. It shows that the periods between 1821-1880, 1911-1930, and 1961-1990 have been the most prolific. The first one, 1821-1880, can be associated with the works of John L. LeConte (724 geadephagan species-group taxa described), Count Dejean (289 taxa), Thomas Say (164 taxa), Baron de Chaudoir (126 taxa), and Victor Motschulsky (121 taxa). The second period, 1911-1930, relates of course to the work of Colonel Casey. By the time of his last publication in 1924, Casey had described 1,864 species-group taxa of carabids based on North American specimens while the number of taxa proposed by all authors combined since Linnaeus amounted to 2,288. The third one, 1961-1990, is connected to the magnificent work of Carl H. Lindroth (76 taxa) on the Carabidae of Canada and Alaska, the work of Thomas Barr (147 taxa) on the cave fauna, and the leadership of George Ball who, besides his own research, directed many students.
Figure 3 shows the growth, through time, of the number of species-group taxa described from North American specimens and those still considered valid. Taking into consideration the trend of description of species-group taxa still valid, the number of native species-group taxa should be slightly over 3,000 by the year 2060.
Table 5 gives statistics regarding the number of geadephagan species-group taxa for each political region covered in this catalogue.
This species, also known as the “Montane Giant Tiger Beetle7,” is restricted to mountains in southeastern Arizona and a small area in western Texas near Big Bend National Park (Pearson et al. 2006: 48).
Excluding the subcosmopolitan , about 105 species (Lorenz 2005: 438-439) in the temperate, subtropical, and tropical areas of the Nearctic (one species), Neotropical (twelve species), Australian (about 17 species), Oriental (about 30 species), eastern Palaearctic (five species), and Afrotropical (about 40 species) Regions. These species are arrayed in 12 subgenera: Perrault (one Neotropical species), Basilewsky (four Afrotropical species), Jeannel (three species in the Philippines, Indonesia, and New Guinea), Perrault9 (two Oriental species), Perrault (three Neotropical species), s.str. (57 species in the Old World as well as the Neotropical and Australian Regions), Jeannel (three Afrotropical species), Jeannel (ten Afrotropical species), (18 species), Jeannel (one Oriental and one Afrotropical species), Jeannel (one Afrotropical species), and Basilewsky (one Afrotropical species).

Sections

"[{\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"T1\"], \"section\": \"Brief history\", \"text\": \"A comparison of the number of valid species and genera between this and previous checklists / catalogues is presented in Table 1.\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"T2\"], \"section\": \"Nomenclatural and distributional information\", \"text\": \"The distributional component consists of a list of state and province records, using the same two-letter postal service style abbreviations used in the 1993 catalogue (Table 2), and a short description of the distribution, usually referring to the northeasternmost, northwesternmost, southwesternmost, and southeasternmost states or provinces. In addition, records for Cape Breton Island, the Queen Charlotte Islands, Vancouver Island, and the Channel Islands are indicated in parentheses after their respective provinces or states. Western Hemisphere countries are listed for species found south of the area covered. States and provinces placed in quotation marks in the descriptive section indicate that only the state or province was given without further precision in the reference cited. The starting point for the distributional records used in this work is Bousquet and Larochelle\\u2019s (1993) catalogue. However, many of their records were undocumented or came from old lists and were not always reliable. State and province records undocumented or considered doubtful are shown in square brackets following the accepted records. Except for the  records which come from identifications generally made by Fritz Hieke, almost all records from CMNH specimens are based on identifications made by Robert L. Davidson, those from LSAM specimens on identifications made by Igor Sokolov, and those from CNC, MCZ, and USNM specimens from identifications or confirmations made by myself. The records provided by Ken Karns and Brian Raber are based on identifications made by Robert L. Davidson.\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"T2\"], \"section\": \"Institution / collection acronyms and abbreviations\", \"text\": \"Besides those used for provinces and states (see Table 2), the following abbreviations are used in the text:\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"FN1\", \"FN2\"], \"section\": \"Nomenclature\", \"text\": \"New taxa. In the xviii and first half of the xix Century it was common practice for authors not to indicate the attribution of the new species-group taxa. Instead, some authors added the word mihi after the specific name, usually to indicate a taxon that the author, himself, was describing. Several collectors provided names for their specimens, even for undescribed ones, and these specimens often circulated among European coleopterists through exchange, gift, or sale. Many undescribed species were subsequently described or illustrated under the collector\\u2019s names by different authors. For these, citations are provided in this catalogue only to the first description or illustration of each species unless the term \\u201cnew species\\u201d or an equivalent expression (such as an asterisk preceding the specific epithet as in Say 1823a1) was included with the species-group name subsequently described or illustrated. Sometimes a species was described / illustrated by different authors the same year under the same names. One example concerned several species (i.e., , , , , and )2 described by Eschscholtz in 1823 in the M\\u00e9moires de la Soci\\u00e9t\\u00e9 Imp\\u00e9riale des Naturalistes de Moscou (volume 6) and illustrated by Fischer von Waldheim on plates available the same year (Sherborn 1922: liii), but included in his Entomographie de la Russie (volume 2) issued in 1824. In such cases, citations are given for the oldest description / illustration (for exceptions see previous entry, Principle of priority) but references to subsequent descriptions / illustrations are noted after the entry of the valid name.\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"FN3\"], \"section\": \"John Lawrence LeConte (1825-1883) Collection\", \"text\": \"LeConte is without doubt the most outstanding North American coleopterist of the xix Century, not only because he described 514 new genus-group and about 4,730 new species-group taxa of beetles (Henshaw 1882: 270), but because he was the first to work seriously on the classification of the North American fauna. During his scientific activity, which lasted almost 40 years, he described 724 new species-group taxa of Geadephaga from North America, 439 (61%) of which were not previously described. LeConte built his collection through his own collecting but also from gifts he received and identifications he provided to many persons from whom he usually retained all or some of the specimens. There is also little doubt that his father, Major John Eatton LeConte3, left his collection to his son. Evidence supporting this can be found in PageBreakLeConte (1856a: 49) when he indicated that his second specimen of  \\u201ccame from the old collection of my father.\\u201d LeConte was a generous man and often offered some of his specimens to visitors (such as Motschulsky [q.v.]) or sent some to acquaintances (such as Chaudoir [q.v.] and Putzeys [q.v.]) though it seems that he retained at least one specimen of each species. Unfortunately in the xix Century the type concept for species-group taxa was not developed and LeConte sometimes gave the only syntype he had in his collection and retained one or more specimens that he acquired after the original descriptions. Therefore, syntypes of some of LeConte\\u2019s species are not in his collection. Moreover, syntypes of some of his species are difficult to find in his collection. LeConte had the habit of mixing the specimens of the new species he considered later as synonyms with those of the valid species. Since many of his specimens only bear a colored disc for label, syntypes of several of his species are not readily ascertainable.\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"FN4\"], \"section\": \"John Lawrence LeConte (1825-1883) Collection\", \"text\": \"In April 1875, LeConte wrote to his friend Alexander Agassiz, director of the Museum of Comparative Zoology in Cambridge, and expressed the wish that his collection be deposited at the museum after his death4. His collection was packed and transported by his longtime friend George Horn. It now stands separate from the general collection along with that of Horn.\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"FN5\"], \"section\": \"Family Carabidae.\", \"text\": \"The current classification of the Carabidae is based mainly on morphological data of adults although molecular sequence data have been used recently to discuss various aspects of carabid phylogeny. Despite several attempts there is no consensus on the classification of several subfamilies or tribes. This is particularly evident among \\u2018basal grade\\u2019 carabids.5\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"T3\"], \"section\": \"Family Carabidae.\", \"text\": \"The world classification of family-group taxa, which has been adopted for the North American fauna in this catalogue, is outlined in Table 3.\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"T4\"], \"section\": \"Adventive species.\", \"text\": \"Table 4 lists the adventive species found on this continent. All but two are accidental introductions (i.e., immigrants). Several carabids were intentionally introduced in New England during the first half of the xx Century for gypsy moth ( Linnaeus) control, including  Kirby,  Linnaeus,  Fabricius,  Herbst,  Linnaeus,  Paykull, and  Linnaeus (see Smith 1959), but only  and  have become established.\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"F1\"], \"section\": \"Adventive species.\", \"text\": \"Most of the remaining 57 adventive species found on this continent prior to the xx Century have probably been introduced with bulk rock, bricks, mortar and no doubt soil taken aboard sailing vessels as ballast and dump ashore on this continent (Brown 1940a; Lindroth 1957c). In the xx Century, species have been introduced into North America likely through shipments of nursery stock, within the soil bagged around the roots, as suggested by Kavanaugh and Erwin (1985), Spence and Spence (1988: 166), and Bousquet (1992a). Despite that quarantine regulations were established in the 1960s for the importation of nursery stock, additional species still continue to land and become established on this continent (Fig. 1). The most recent one is  in the Willamette Valley in northwestern Oregon (Kavanaugh and LaBonte 2008).\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"F1\"], \"section\": \"Adventive species.\", \"text\": \"Figure 1 illustrates the arrival of adventive species in North America through time based on the collecting date of the first inventoried specimen. The rate of arrival has been steady in the xx Century.\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"F2\"], \"section\": \"Native species.\", \"text\": \"Figure 2 illustrates through decades the number of North American species-group taxa described and the number of those regarded as valid today based on the present catalogue. It shows that the periods between 1821-1880, 1911-1930, and 1961-1990 have been the most prolific. The first one, 1821-1880, can be associated with the works of John L. LeConte (724 geadephagan species-group taxa described), Count Dejean (289 taxa), Thomas Say (164 taxa), Baron de Chaudoir (126 taxa), and Victor Motschulsky (121 taxa). The second period, 1911-1930, relates of course to the work of Colonel Casey. By the time of his last publication in 1924, Casey had described 1,864 species-group taxa of carabids based on North American specimens while the number of taxa proposed by all authors combined since Linnaeus amounted to 2,288. The third one, 1961-1990, is connected to the magnificent work of Carl H. Lindroth (76 taxa) on the Carabidae of Canada and Alaska, the work of Thomas Barr (147 taxa) on the cave fauna, and the leadership of George Ball who, besides his own research, directed many students.\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"F3\"], \"section\": \"Native species.\", \"text\": \"Figure 3 shows the growth, through time, of the number of species-group taxa described from North American specimens and those still considered valid. Taking into consideration the trend of description of species-group taxa still valid, the number of native species-group taxa should be slightly over 3,000 by the year 2060.\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"T5\"], \"section\": \"Political regions.\", \"text\": \"Table 5 gives statistics regarding the number of geadephagan species-group taxa for each political region covered in this catalogue.\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"FN7\"], \"section\": \"Distribution.\", \"text\": \"This species, also known as the \\u201cMontane Giant Tiger Beetle7,\\u201d is restricted to mountains in southeastern Arizona and a small area in western Texas near Big Bend National Park (Pearson et al. 2006: 48).\"}, {\"pmc\": \"PMC3577090\", \"pmid\": \"23431087\", \"reference_ids\": [\"FN9\"], \"section\": \"Diversity.\", \"text\": \"Excluding the subcosmopolitan , about 105 species (Lorenz 2005: 438-439) in the temperate, subtropical, and tropical areas of the Nearctic (one species), Neotropical (twelve species), Australian (about 17 species), Oriental (about 30 species), eastern Palaearctic (five species), and Afrotropical (about 40 species) Regions. These species are arrayed in 12 subgenera:  Perrault (one Neotropical species),  Basilewsky (four Afrotropical species),  Jeannel (three species in the Philippines, Indonesia, and New Guinea),  Perrault9 (two Oriental species),  Perrault (three Neotropical species),  s.str. (57 species in the Old World as well as the Neotropical and Australian Regions),  Jeannel (three Afrotropical species),  Jeannel (ten Afrotropical species),  (18 species),  Jeannel (one Oriental and one Afrotropical species),  Jeannel (one Afrotropical species), and  Basilewsky (one Afrotropical species).\"}]"

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