Harpalinae is a huge

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...

ground beetle Ground beetles are a large, cosmopolitan distribution, cosmopolitan family (biology), family of beetles, the Carabidae, with more than 40,000 species worldwide, around 2,000 of which are found in North America and 2,700 in Europe. As of 2015, it i ...

s that contains 20,000 species or ~6,400 spp. in 24 tribes worldwide, according to others. A rarely used common name for the subfamily is the harp beetles. The Harpalinae contain the most

apomorph In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...

ic ground beetles, displaying a wide range of forms and behaviors. Some are, rare among ground beetles,

omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutr ...

s or even

herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...

s. Many closely related subfamilies have been treated as subordinate

taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

of the Harpalinae by various authors. Among these are the

Dryptinae Dryptinae is a subfamily of beetles in the family Carabidae, containing the following genera: * '' Acrogenys'' Macleay, 1864 * '' Agastus'' Schmidt-Goebel, 1846 * '' Ancystroglossus'' Chaudoir, 1862 * '' Chaudoirella'' Mateu, 1982 * '' Coarazuph ...

Lebiinae Lebiinae is a subfamily of beetles in the family Carabidae. Genera The subfamily includes the following genera: * '' Abrodiella'' Bousquet, 2002 * '' Actenonyx'' White, 1846 * '' Aeolodermus'' Andrewes, 1929 * '' Afrodromius'' Basilewsky, 1958 ...

(including Cyclosominae, Mormolycinae, Odacanthinae, Perigoninae),

Licininae Licininae is a subfamily of beetles in the family Carabidae, containing the following genera: * '' Acanthoodes'' Basilewsky, 1953 * '' Actodus'' Alluaud, 1915 * '' Acutosternus'' Lecordier & Girard, 1988 * '' Adelopomorpha'' Heller, 1916 * '' An ...

(including Chlaeniinae, Oodinae),

Orthogoniinae Orthogoniinae is a subfamily of ground beetles (family Carabidae). Occasionally it was treated as a tribe Orthogoniini of subfamily Harpalinae Harpalinae is a huge subfamily of ground beetles that contains 20,000 species or ~6,400 spp. in 24 ...

, Panagaeinae,

Platyninae Platyninae is a subfamily of ground beetles (family Carabidae). Genera The subfamily Platyninae contains about 250 genera organized into three tribes: ; Tribe Omphreini Ganglbauer, 1891 : '' Omphreus'' Dejean, 1828 ; Tribe Platynini Bonelli, ...

Pseudomorphinae Pseudomorphinae is a subfamily of ground beetles (family Carabidae). Occasionally it was treated as a tribe Pseudomorphini of subfamily Harpalinae, particularly when this was circumscribed loosely. It contains the following genera: * ''Adelotop ...

Pterostichinae Pterostichinae is a subfamily of ground beetles (family Carabidae). It belongs to the advanced harpaline assemblage, and if these are circumscribed ''sensu lato'' as a single subfamily, Pterostichinae are downranked to a tribe Pterostichini. How ...

(including Zabrinae). Here, they are considered independent families within the harpaline (''

sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...

'') assemblage, and this is also tentatively assumed for the enigmatic

monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

Ginema ''Ginema'' is a genus of ground beetles (family Carabidae). The single described species, ''Ginema thomasi'', is known only from Bolivia. Its closest living relatives are unknown, and is sometimes separated as subfamily Gineminae. While this spe ...

''.

Systematics

At least 4 large and several smaller supertribes can be distinguished. The former " Pterostichitae" on the other hand seem to be, as presently delimited, a

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

assemblage, but probably also contains a number of

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...

s closer to each other than to other Harpalinae. The

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

around the

type genus In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal f ...

Pterostichus ''Pterostichus'' is a very large genus of ground beetles with a Holarctic distribution in the subfamily Harpalinae. It has over 1,100 species. The beetles are predatory, but sometimes feed on strawberries. They can be found under rocks and prefer ...

'' is thus recognized as subfamily

here.

Genera

The Harpalinae include the following genera: Harpalus affinis01

* '' Acinopus'' Dejean, 1821 * '' Acupalpus'' Latreille, 1829 * '' Afromizonus'' Basilewsky, 1947 * '' Agonidus'' Casey, 1914 * '' Allendia'' Noonan, 1974 * '' Allocinopus'' Broun, 1903 * '' Allosiopelus'' Ito, 1995 * '' Amblygnathus'' Dejean, 1829 * '' Amblystomus'' Erichson, 1837 * '' Amerinus'' Casey, 1884 * '' Amphasia'' Newman, 1836 * '' Angionychus'' Klug, 1853 * '' Anisocnemus'' Chaudoir, 1843 * '' Anisodactylus'' Dejean, 1829 * '' Anisostichus'' Emden, 1953 * '' Anomostomus'' Laferte-Senectere, 1853 * '' Anthracus''

Motschulsky Victor Ivanovich Motschulsky (sometimes Victor von Motschulsky, russian: link=no, Виктор Иванович Мочульский, 11 April 1810, in St. Petersburg – 5 June 1871, in Simferopol) was a Russian entomologist mainly interested in ...

, 1850 * ''

Athrostictus ''Athrostictus'' is a genus of ground beetles in the family Carabidae. Species These 18 species belong to the genus ''Athrostictus'': * '' Athrostictus chlaenioides'' (Dejean, 1829) * '' Athrostictus circumfusus'' (Putzeys, 1878) * '' Athrosti ...

'' Bates, 1878 * '' Axinotoma'' Dejean, 1829 * '' Aztecarpalus'' Ball, 1970 * '' Barysomus'' Dejean, 1829 * '' Batoscelis'' Dejean, 1836 * '' Bleusea'' Bedel, 1897 * '' Boeomimetes'' Peringuey, 1896 * '' Bottchrus'' Jedlicka, 1935 * '' Bradybaenus'' Dejean, 1829 * '' Bradycellus'' Erichson, 1837 * '' Bradycidus'' Casey, 1914 * '' Bronislavia'' Semenov, 1891 * '' Carenochyrus'' Solsky, 1874 * '' Carterus'' Dejean, 1830 * '' Cenogmus'' Sloane, 1898 * '' Chilotomus'' Chaudoir, 1842 * '' Chydaeus'' Chaudoir, 1854 * '' Coleolissus'' Bates, 1892 * '' Crasodactylus'' Guerin-Meneville, 1847 * '' Cratacanthus'' Dejean, 1829 * '' Cratognathus'' Dejean, 1829 * '' Cratosoma'' Jeannel, 1948 * '' Criniventer'' van Emdem, 1953 * '' Cyptomicrus'' Vinson, 1939 * '' Daptus'' Fischer von Waldheim, 1823 * '' Diachromus'' Erichson, 1837 * ''

Dicheirotrichus ''Dicheirotrichus'' is a genus of beetles in the family Carabidae Ground beetles are a large, cosmopolitan family of beetles, the Carabidae, with more than 40,000 species worldwide, around 2,000 of which are found in North America and 2,700 i ...

'' Jacqelin du Val, 1857 (including ''Trichocellus'') * '' Dicheirus'' Mannerheim, 1843 * '' Dioryche'' W.S. MacLeay, 1825 * '' Discoderus'' LeConte, 1853 * '' Ditomus'' Bonelli, 1810 * '' Dixus'' Billberg, 1820 * '' Dregus'' Motscheisky, 1864 * '' Ectinothorax'' Alluaud, 1941 * '' Egadyla'' Alluaud, 1916 * '' Eocarterus'' Stichel, 1923 * '' Eriophonus'' Tschitscherine, 1901 * '' Eucarterus'' Reitter, 1900 * '' Euryderus'' LeConte, 1846 * '' Euthenarus'' Bates, 1874 * '' Gaioxenus'' Broun 1910 * '' Geodromus'' Dejean, 1829 * '' Geopinus'' LeConte, 1848 * '' Gnathaphanus'' W.S. MacLeay, 1825 * '' Goniocellus'' Casey, 1914 * '' Graniger'' Motschulsky, 1864 * '' Gugheorites'' Basilewsky, 1951 * '' Gynandromorphus'' Dejean, 1829 * '' Hakaharpalus'' Larochelle & Lariviere, 2005 * '' Haplaner'' Chaudoir, 1878 * '' Haplanister'' Moore, 1996 * '' Harpalinus'' Jeannel, 1946 * '' Harpalobrachys'' Tschitscherine, 1899 * '' Harpalodiodes'' Bousquet, 2002 * '' Harpalomimetes'' Schauberger, 1933 * '' Harpalomorphus'' Peringuey, 1896 * ''

Harpalus Harpalus (Greek: Ἅρπαλος) son of Machatas was an aristocrat of Macedon and boyhood friend of Alexander the Great in the 4th century BC. Harpalus was repeatedly entrusted with official roles by Alexander and absconded three times with la ...

'' Latreille, 1802 (including ''Harpalobrachys'') * '' Harpathaumas'' Basilewsky, 1947 * '' Harponixus'' Basilewsky, 1950 * '' Hartonymus'' Casey, 1914 * '' Hemiaulax'' Bates, 1892 * '' Heteracantha'' Brulle, 1834 * '' Hippoloetis'' Laporte, 1835 * '' Hyphaereon'' W.S. MacLeay, 1825 * '' Hypharpax'' W.S. McLeay, 1825 * '' Idiomelas'' Tschitscherine, 1900 * '' Indiophonus'' N.Ito, 1996 * '' Kaffovatus'' Clarke, 1972 * '' Kenyacus'' Alluaud, 1917 * '' Kiwiharpalus'' Larochelle & Lariviere, 2005 * '' Kupeharpalus'' Larochelle & Lariviere, 2005 * '' Lecanomerus'' Chaudoir, 1850 * '' Liochirus'' Tschitscherine, 1897 * '' Liodaptus'' Bates, 1889 * '' Lioholus'' Tschitscherine, 1897 * '' Loxoncus'' Schmidt-Gobel, 1846 * '' Machozetus'' Chaudoir, 1850 * '' Maoriharpalus'' Larochelle & Lariviere, 2005 * ''

Meroctenus ''Meroctenus'' is a genus of beetle in the family Carabidae first described by Max Gemminger, Gemminger & Edgar von Harold, Harold in 1868. Species ''Meroctenus'' contains the following ten species: * ''Meroctenus crenulatus'' (Chaudoir, 1843 ...

'' Gemminger et Harold, 1868 * '' Microderes'' Faldermann, 1836 * '' Nemaglossa'' Solier, 1849 * '' Neoaulacoryssus'' Noonan, 1985 * '' Neodiachipteryx'' Noonan, 1985 * '' Neohyparpalus'' Clarke, 1981 * '' Neophygas'' Noonan, 1976 * '' Nesarpalus'' Bedel, 1897 * '' Nipponoharpalus'' Habu, 1973 * '' Nornalupia'' Kataev, 2002 * '' Nothodaptus'' Maindron, 1906 * '' Notiobia'' Perty, 1830 * '' Notospeophonus'' Moore, 1962 * '' Odontocarus'' Solier, 1835 * '' Oedesis'' Motschulsky, 1850 * '' Oesyperus'' Andrewes, 1923 * '' Omostropus'' Peringuey, 1896 * '' Ooidius'' Chaudoir, 1847 * '' Oosoma'' Nietner, 1857 * '' Ophoniscus'' Bates, 1892 * '' Ophonus'' Dejean, 1821 * '' Orthogenium'' Chaudoir, 1835 * '' Oxycentrus'' Chaudoir, 1854 * '' Pachycarus'' Solier, 1835 * '' Pachytrachelus'' Chaudoir, 1852 * '' Panagrius'' Andrewes, 1933 * '' Pangus'' Dejean, 1821 * '' Parabaris'' Broun, 1881 * '' Paramecus'' Dejean, 1829 * '' Parasiopelus'' Basilewsky, 1950 * '' Paregaploa'' Muller, 1947 * '' Parophonus'' Ganglbauer, 891/small> * '' Paulianoscirtus'' Basilewsky, 1976 * '' Pelmatellus'' Bates, 1882 * '' Penthophonus'' Reitter, 1900 * ''

Penthus In Greek mythology, Penthus (Πενθος) was the personification of grief. When Zeus Zeus or , , ; grc, Δῐός, ''Diós'', label=Genitive case, genitive Aeolic Greek, Boeotian Aeolic and Doric Greek#Laconian, Laconian grc-dor, Δ� ...

'' Chaudoir, 1843 * '' Phanagnathus'' Basilewsky, 1950 * '' Phobophorus'' Motschulsky, 1850 * '' Pholeodytes'' Britton, 1962 * '' Phorticosomus'' Shaum, 1863 * '' Phyrometus'' Basilewsky, 1946 * '' Piosoma'' LeConte, 1848 * '' Platymetopsis'' Ball et Maddison, 1987 * '' Platymetopus'' Dejean, 1829 * '' Pogonodaptus'' Horn, 1881 * '' Polpochila'' Solier, 1849 * ''

Prakasha Prakasha, popularly known as Dakshin Kashiis a village situated on the bank of Tapi River in taluka Shahada in Nandurbar district, Maharashtra, India. Demographics Prakasha has a population of around 20,000, of which 90% is in agriculture a ...

'' Andrewes, 1919 * '' Proditomus'' Schauberger, 1934 * '' Progonochaetus'' J. Muller, 1938 * '' Pseudanisotarsus'' Noonan, 1973 * '' Pseudaristus'' Reitter, 1900 * '' Pseudodiachipteryx'' Burgeon, 1936 * '' Pseudognathaphanus'' Schauberger, 1932 * '' Pseudohyparpalus'' Basilewsky, 1946 * '' Pseudorhysopus'' Kataev et Wrase, 2001 * '' Pseudoselenophorus'' Peringuey, 1896 * '' Psychristus'' Andrewes, 1930 * '' Rhabidius'' Basilewsky, 1948 * '' Rhysopus'' Andrewes, 1929 * '' Scybalicus'' Schaum, 1862 * '' Selenophorus'' Dejean, 1831 * '' Selenotichnus'' Kataev, 1999 * '' Siopelus'' Murray, 1859 * '' Stenolophidius'' Jeannel, 1948 * '' Stenolophus'' Dejean, 1821 * '' Stenomorphus'' Dejean, 1831 * '' Syllectus'' Bates, 1878 * '' Trachysarus'' Reed, 1874 * '' Trichopselaphus'' Chaudoir, 1843 * '' Trichotichnus'' A.Morawitz, 1863 * '' Trichoxycentrus'' N. Ito, 2000 * '' Triplosarus'' Bates, 1874 * '' Tropicoritus'' Alluaud, 1917 * '' Tschitscherinellus'' Csiki, 1906 * '' Tuiharpalus'' Larochelle & Lariviere, 2005 * '' Typsiharpalus'' Tschitscherine, 1901 * '' Uenanthracus'' Kasahara, 1994 * '' Veradia'' Castelnau, 1867 * '' Xenodochus'' Andrewes, 1941 * '' Xenophonus'' Muller, 1942 * '' Xestonotus'' LeConte, 1853

Fossil genera

* †'' Burmapseudomorphus'' Beutel et al. 2020

Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...

, Myanmar,

Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...

, Tribe Pseudomorphini

References

* (2008)
Trees of family Carabidae
. Retrieved 2008-JUL-24. {{Taxonbar, from=Q137373 Carabidae subfamilies Taxa named by Franco Andrea Bonelli