Ground beetles are a large,
cosmopolitan
Cosmopolitan may refer to:
Food and drink
* Cosmopolitan (cocktail), also known as a "Cosmo"
History
* Rootless cosmopolitan, a Soviet derogatory epithet during Joseph Stalin's anti-Semitic campaign of 1949–1953
Hotels and resorts
* Cosmopoli ...
family
Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
of
beetle
Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s,
the Carabidae, with more than 40,000 species worldwide, around 2,000 of which are found in
North America and 2,700 in
Europe
Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...
. As of 2015, it is one of the 10 most species-rich animal families. They belong to the
Adephaga. Members of the family are primarily carnivorous, but some members are
phytophagous
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 mouthpar ...
or omnivorous.
Description and ecology
Although their body shapes and coloring vary somewhat, most are shiny black or metallic and have ridged wing covers (
elytra
An elytron (; ; , ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometimes alterna ...
). The elytra are fused in some
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, particularly the large
Carabinae
Carabinae is a subfamily of beetles in the family Carabidae, containing the following genera:
* '' Aplothorax'' Waterhouse, 1841
* ''Calosoma'' Weber, 1801
*'' Calosoma oregonus'' Gidaspow, 1959
* ''Carabus'' Linnaeus, 1758
* '' Ceroglossus'' So ...
, rendering the beetles unable to fly. The
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
''
Mormolyce phyllodes'' is known as violin beetle due to their peculiarly shaped elytra. All carabids except the quite primitive
flanged bombardier beetles (Paussinae) have a groove on their
fore leg tibiae bearing a comb of hairs used for cleaning their
antennae.
Defensive secretions
Typical for the ancient beetle
suborder Adephaga to which they belong, they have paired
pygidial
The pygidium (plural pygidia) is the posterior body part or shield of crustaceans and some other arthropods, such as insects and the extinct trilobites. In groups other than insects, it contains the anus and, in females, the ovipositor. It is com ...
glands in the lower back of the
abdomen
The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
. These are well developed in ground beetles, and produce noxious or even
caustic
Caustic most commonly refers to:
* Causticity, a property of various corrosive substances
** Sodium hydroxide, sometimes called ''caustic soda''
** Potassium hydroxide, sometimes called ''caustic potash''
** Calcium oxide, sometimes called ''caus ...
secretions used to deter would-be
predator
Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s. In some, commonly known as
bombardier beetles, these secretions are mixed with
volatile compounds and ejected by a small
combustion
Combustion, or burning, is a high-temperature exothermic redox chemical reaction between a fuel (the reductant) and an oxidant, usually atmospheric oxygen, that produces oxidized, often gaseous products, in a mixture termed as smoke. Combus ...
, producing a loud popping sound and a cloud of hot and acrid gas that can injure small
mammals, such as
shrew
Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to differ ...
s, and is liable to kill
invertebrate
Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
predators outright.
To humans, getting "bombed" by a bombardier beetle is a decidedly unpleasant experience. This ability has
evolved
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
independently twice, as it seems, in the
flanged bombardier beetles (Paussinae), which are among the most ancient ground beetles, and in the typical bombardier beetles (
Brachininae
Brachininae is a subfamily 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 in Europ ...
), which are part of a more "modern" lineage. The
Anthiini, though, can mechanically squirt their defensive secretions for considerable distances and are able to aim with a startling degree of accuracy; in
Afrikaans
Afrikaans (, ) is a West Germanic language that evolved in the Dutch Cape Colony from the Dutch vernacular of Holland proper (i.e., the Hollandic dialect) used by Dutch, French, and German settlers and their enslaved people. Afrikaans gra ...
, they are known as ' ("eye-pissers"). In one of the very few known cases of a
vertebrate
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with c ...
mimicking an
arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
, juvenile ''
Heliobolus lugubris''
lizards are similar in color to the
aposematic
Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste o ...
''oogpister'' beetles, and move in a way that makes them look surprisingly similar to the insects at a casual glance.
A folk story claims that
Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
once found himself on the receiving end of a bombardier beetle's attack, based on a passage in his autobiography.
Darwin stated in a letter to
Leonard Jenyns
Leonard Jenyns (25 May 1800 – 1 September 1893) was an English clergyman, author and naturalist. He was forced to take on the name Leonard Blomefield to receive an inheritance. He is chiefly remembered for his detailed phenology observations ...
that a beetle had attacked him on that occasion, but he did not know what kind:
A '' Cychrus rostratus'' once squirted into my eye & gave me extreme pain; & I must tell you what happened to me on the banks of the Cam
Calmodulin (CaM) (an abbreviation for calcium-modulated protein) is a multifunctional intermediate calcium-binding messenger protein expressed in all eukaryotic cells. It is an intracellular target of the secondary messenger Ca2+, and the bin ...
in my early entomological days; under a piece of bark I found two carabi (I forget which) & caught one in each hand, when lo & behold I saw a sacred '' Panagæus crux major''; I could not bear to give up either of my Carabi, & to lose ''Panagæus'' was out of the question, so that in despair I gently seized one of the carabi between my teeth, when to my unspeakable disgust & pain the little inconsiderate beast squirted his acid down my throat & I lost both Carabi & ''Panagæus''!
Ecology
Common
habitat
In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s are under the bark of trees, under logs, or among rocks
or sand by the edge of ponds and rivers. Most species are
carnivorous and actively hunt for any
invertebrate
Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
prey they can overpower.
Some run swiftly to catch their prey;
tiger beetle
Tiger beetles are a family of beetles, Cicindelidae, known for their aggressive predatory habits and running speed. The fastest known species of tiger beetle, ''Rivacindela hudsoni'', can run at a speed of , or about 125 body lengths per second. ...
s (Cicindelinae) can sustain speeds of
– in relation to their body length they are among the fastest land animals on Earth. Unlike most Carabidae, which are
nocturnal, the tiger beetles are active diurnal hunters and often brightly coloured; they have large eyes and hunt by sight. Ground beetles of the species ''
Promecognathus laevissimus'' are specialised predators of the cyanide millipede ''
Harpaphe haydeniana'', countering the
hydrogen cyanide that makes these millipedes poisonous to most carnivores.
Relationship with humans
As predators of invertebrates, including many
pests, most ground beetles are considered beneficial organisms. The
caterpillar hunter
''Calosoma'' is a genus of large ground beetles that occur primarily throughout the Northern Hemisphere, and are referred to as caterpillar hunters or caterpillar searchers. Many of the 167 species are largely or entirely black, but some have ...
s (''Calosoma'') are famous for their habit of devouring prey in quantity, eagerly feeding on
tussock moth
The Lymantriinae (formerly called the Lymantriidae) are a subfamily of moths of the family Erebidae. The taxon was erected by George Hampson in 1893.
Many of its component species are referred to as "tussock moths" of one sort or another. The cat ...
(Lymantriidae) caterpillars,
processionary caterpillar
Thaumetopoeinae is a subfamily of moths in the family Notodontidae. This group is sometimes treated as a family (biology), family Thaumetopoeidae with three subfamilies: Thaumetopoeinae, Anaphinae and Epicominae. However, it is now commonly treat ...
s (Thaumetopoeidae) and
woolly worms (Arctiidae), which, due to their
urticating hair
Urticating hairs or urticating bristles are one of the primary defense mechanisms used by numerous plants, almost all New World tarantulas, and various lepidopteran caterpillars. ''Urtica'' is Latin for "nettle" (stinging nettles are in the genu ...
s, are avoided by most insectivores. Large numbers of the
forest caterpillar hunter (''C. sycophanta''), native to
Europe
Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...
, were shipped to
New England
New England is a region comprising six states in the Northeastern United States: Connecticut, Maine, Massachusetts, New Hampshire, Rhode Island, and Vermont. It is bordered by the state of New York (state), New York to the west and by the Can ...
for
biological control
Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also i ...
of the
gypsy moth
''Lymantria dispar'', also known as the gypsy moth or the spongy moth, is a species of moth in the family Erebidae. ''Lymantria dispar'' is subdivided into several subspecies, with subspecies such as ''L. d. dispar'' and ''L. d. japonica'' bei ...
(''Lymantria dispar'') as early as 1905.
A few species are nuisance pests. ''
Zabrus'' is one of the few
herbivorous
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 mouthpar ...
ground beetle genera, and on rare occasions ''
Zabrus tenebrioides'', for example, occurs abundantly enough to cause some damage to
grain crops. Large species, usually the
Carabinae
Carabinae is a subfamily of beetles in the family Carabidae, containing the following genera:
* '' Aplothorax'' Waterhouse, 1841
* ''Calosoma'' Weber, 1801
*'' Calosoma oregonus'' Gidaspow, 1959
* ''Carabus'' Linnaeus, 1758
* '' Ceroglossus'' So ...
, can become a nuisance if present in large numbers, particularly during outdoor activities such as
camping
Camping is an outdoor activity involving overnight stays away from home, either without shelter or using basic shelter such as a tent, or a recreational vehicle. Typically, participants leave developed areas to spend time outdoors in more na ...
; they void their defensive secretions when threatened, and in hiding among provisions, their presence may spoil food. Since ground beetles are generally reluctant or even unable to fly, mechanically blocking their potential routes of entry is usually easy. The use of insecticides specifically for carabid intrusion may lead to unfortunate side effects, such as the release of their secretions, so it generally is not a good idea unless the same applications are intended to exclude ants, parasites or other crawling pests.
![Panagaeus cruxmajor bl](https://upload.wikimedia.org/wikipedia/commons/7/7a/Panagaeus_cruxmajor_bl.jpg)
Especially in the 19th century and to a lesser extent today, their large size and conspicuous coloration, as well as the odd
morphology
Morphology, from the Greek and meaning "study of shape", may refer to:
Disciplines
* Morphology (archaeology), study of the shapes or forms of artifacts
* Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
of some (e.g. the
Lebiini), made many ground beetles a popular object of collection and study for professional and amateur
coleopterologists. High prices were paid for rare and exotic specimens, and in the early to mid-19th century, a veritable "beetle craze" occurred in England. As mentioned above, Charles Darwin was an ardent collector of beetles when he was about 20 years old, to the extent that he would rather scour the countryside for rare specimens with
William Darwin Fox
The Reverend William Darwin Fox (23 April 1805 – 8 April 1880) was an English clergyman, naturalist, and a second cousin of Charles Darwin.
Early life
Fox was born in 1805 and initially raised at Thurleston Grange near Elvaston, Derbys ...
,
John Stevens Henslow
John Stevens Henslow (6 February 1796 – 16 May 1861) was a British priest, botanist and geologist. He is best remembered as friend and mentor to his pupil Charles Darwin.
Early life
Henslow was born at Rochester, Kent, the son of a solicit ...
, and
Henry Thompson than to study
theology
Theology is the systematic study of the nature of the divine and, more broadly, of religious belief. It is taught as an academic discipline, typically in universities and seminaries. It occupies itself with the unique content of analyzing the ...
as his father wanted him to do. In his autobiography, he fondly recalled his experiences with ''
Licinus'' and ''
Panagaeus'', and wrote:
No poet ever felt more delight at seeing his first poem published than I did at seeing in Stephen's ''Illustrations of British Insects'' the magic words, "captured by C. Darwin, Esq."
Evolution and systematics
The
Adephaga are documented since the end of the
Permian
The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last ...
, about (Mya). Ground beetles
evolved
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
in the latter
Triassic
The Triassic ( ) is a geologic period and system (stratigraphy), system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Year#Abbreviations yr and ya, Mya), to the beginning of the Jurassic Period 201.36 ...
, having separated from their closest relatives by 200 Mya. The family diversified throughout the
Jurassic
The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
, and the more advanced lineages, such as the
Harpalinae
Harpalinae is a huge subfamily of ground beetles 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 apomorphic ...
, underwent a vigorous radiation starting in the
Cretaceous
The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
. The closest living relatives of the ground beetles are the
false ground beetle
The Trachypachidae (sometimes known as false ground beetles) are a family of beetles that generally resemble small ground beetles, but that are distinguished by the large coxae of their rearmost legs. There are only six known extant species in th ...
s (Trachypachidae) and the
tiger beetle
Tiger beetles are a family of beetles, Cicindelidae, known for their aggressive predatory habits and running speed. The fastest known species of tiger beetle, ''Rivacindela hudsoni'', can run at a speed of , or about 125 body lengths per second. ...
s (Cicindelidae). They are sometimes even included in the Carabidae as subfamilies or as tribes ''
incertae sedis
' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
'', but more preferably they are united with the ground beetles in the
superfamily Caraboidea, or
Geadephaga.
[Vasilikopoulos, A., Balke, M., Kukowka, S., Pflug, J.M., Martin, S., Meusemann, K., Hendrich, L., Mayer, C., Maddison, D.R., Niehuis, O., Beutel, R.G. and Misof, B. (2021), Phylogenomic analyses clarify the pattern of evolution of Adephaga (Coleoptera) and highlight phylogenetic artefacts due to model misspecification and excessive data trimming. Syst Entomol, 46: 991-1018. https://doi.org/10.1111/syen.12508]
Much research has been done on elucidating the
phylogeny
A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spe ...
of the ground beetles and adjusting
systematics and
taxonomy
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
accordingly. While no completely firm consensus exists, a few points are generally accepted: The ground beetles seemingly consist of a number of more
basal lineages and the extremely diverse Harpalinae, which contain over half the described species and into which several formerly independent families had to be subsumed.
Subfamilies and selected genera
The taxonomy used here is primarily based on the ''Catalogue of Palaearctic Coleoptera'' and the ''Carabidae of the World Database''. Other classifications, while generally agreeing with the division into a basal radiation of more primitive lineages and the more advanced group informally called "Carabidae Conjunctae", differ in details. For example, the system used by the
Tree of Life Web Project
The Tree of Life Web Project is an Internet project providing information about the diversity and phylogeny of life on Earth.
This collaborative peer reviewed project began in 1995, and is written by biologists from around the world. The site ...
makes little use of subfamilies, listing most
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 is in the discipline of anthropology. This definition is contested, in part due to confli ...
s as ''
incertae sedis
' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
'' as to subfamily. ''
Fauna Europaea Fauna Europaea is a database of the scientific names and distribution of all living multicellular Europe
Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight o ...
'', though,
splits rather than lumps the Harpalinae, restricting them to what in the system used here is the tribe
Harpalini
Harpalini is a tribe of a diverse group of ground beetles belonging to the subfamily Harpalinae within the broader family Carabidae. The tribe contains more than 1,900 species.
Genera
These 103 genera belong to Harpalini:
; Subtribe Amblystom ...
.
The exclusion of Trachypachidae and Cicindelidae as separate families is now amply supported, as is the inclusion of Rhysodidae as a subfamily, closely related to
Paussinae and
Siagoninae
Siagoninae is a subfamily of beetles in the family Carabidae. It contains 83 species in 3 genera:
* Genus ''Enceladus(insect), Enceladus'' Bonelli, 1813
** ''Enceladus gigas'' Bonelli, 1813
* Genus ''Luperca'' Castelnau, 1840
** ''Luperca goryi' ...
.
[
The inclusive Harpalinae presented here are used for two reasons, one scientific and one practical – first, the majority of authors presently use this system, following the ''Catalogue of Palaearctic Coleoptera''. Second, the ]MediaWiki
MediaWiki is a free and open-source wiki software. It is used on Wikipedia and almost all other Wikimedia websites, including Wiktionary, Wikimedia Commons and Wikidata; these sites define a large part of the requirement set for MediaWi ...
markup cannot at present adequately represent the relationships of the ground beetle subgroups in detail if the restricted view of the Harpalinae is chosen.
Basal ground beetles
Carabinae
Carabinae is a subfamily of beetles in the family Carabidae, containing the following genera:
* '' Aplothorax'' Waterhouse, 1841
* ''Calosoma'' Weber, 1801
*'' Calosoma oregonus'' Gidaspow, 1959
* ''Carabus'' Linnaeus, 1758
* '' Ceroglossus'' So ...
Latreille, 1802 – including Agoninae and Callistinae
* '' Altagonum''
* ''Aplothorax
''Aplothorax burchelli'' is a species of beetle in the family Carabidae, the only species in the genus ''Aplothorax''. It is endemic to the island of Saint Helena in the South Atlantic. The genus has been found to be nested within ''Calosoma
...
'' (monotypic genus)
* ''Calosoma
''Calosoma'' is a genus of large ground beetles that occur primarily throughout the Northern Hemisphere, and are referred to as caterpillar hunters or caterpillar searchers. Many of the 167 species are largely or entirely black, but some have ...
'' - including ''Callisthenes''
* ''Carabus
''Carabus'' is a genus of beetles in family Carabidae. The genus is highly diverse with 94 subgenera, 897 species and 2300 subspecies, thus is the largest genus in the subfamily Carabinae.Deuve T.; Cruaud, A.; Genson, G.; and Rasplus, J.Y. (201 ...
''
* ''Cychrus
''Cychrus'' is a large genus of rare snail-eating beetles in the family Carabidae. There are at least 160 described species in ''Cychrus''. They are found throughout the world, although more than 80 percent of the species occur in China.
''Cychr ...
''
* ''Homothes
''Homethes'' is a genus in the beetle family Carabidae. There are about 10 described species in ''Homethes''.
Species
These 10 species belong to the genus ''Homethes'':
* ''Homethes angulatus'' Blackburn, 1892 (Australia)
* ''Homethes elegans'' ...
''
* ''Fortagonum
''Fortagonum'' is a genus in the beetle family Carabidae. There are more than 20 described species in ''Fortagonum'', found in Indonesia and New Guinea.
Species
These 29 species belong to the genus ''Fortagonum'':
* ''Fortagonum acuticolle'' Ba ...
''
* '' Laemostenus''
* '' Notagonum''
Cicindinae
Elaphrinae Latreille, 1802
* '' Blethisa''
* '' Diacheila'' 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 ...
, 1844
* ''Elaphrus
''Elaphrus'' is a genus of ground beetle native to the Palearctic, the Nearctic, the Near East and Northern Africa
North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singula ...
''
Hiletinae
Loricerinae
Loricerinae is a subfamily 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 in Europ ...
Bonelli, 1810
* '' Loricera''
Migadopinae
Nebriinae (includes Notiophilinae, often included in Carabinae)
* '' Leistus''
* '' Nebria''
* ''Notiophilus
''Notiophilus'' is a genus of ground beetle native to the Palearctic, the Nearctic, the Near East and North Africa. It contains the following 57 species:
*'' Notiophilus aeneus'' (Herbst, 1806)
*'' Notiophilus aestuans'' Dejean, 1826
*'' Notioph ...
''
* '' Pelophila'' Dejean, 1821
* '' Nippononebria''
Nototylinae
Nototylinae is a subfamily 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 in Euro ...
Omophroninae Bonelli, 1810 – round sand beetles
* '' Omophron''
Paussinae – ant nest beetles, flanged bombardier beetles
Promecognathinae
Promecognathinae is a subfamily 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 in ...
Rhysodinae
Rhysodinae is a subfamily (sometimes called wrinkled bark beetles) in the family Carabidae. There are 19 genera and at least 380 described species in Rhysodinae. The group of genera making up Rhysodinae had been treated as the family Rhysodidae i ...
– wrinkled bark beetles
Scaritinae Bonelli, 1810 – pedunculate ground beetles
* '' Clivina''
* '' Dyschirius'' Bonelli, 1810
* '' Scarites''
Siagoninae
Siagoninae is a subfamily of beetles in the family Carabidae. It contains 83 species in 3 genera:
* Genus ''Enceladus(insect), Enceladus'' Bonelli, 1813
** ''Enceladus gigas'' Bonelli, 1813
* Genus ''Luperca'' Castelnau, 1840
** ''Luperca goryi' ...
Bonelli, 1810
Carabidae Conjunctae
Amblytelinae Sloane, 1898
* '' Amblytelus''
Apotominae
Brachininae
Brachininae is a subfamily 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 in Europ ...
– typical bombardier beetles
Broscinae
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 ...
(sometimes in Harpalinae)
Gineminae (sometimes in Harpalinae)
Harpalinae
Harpalinae is a huge subfamily of ground beetles 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 apomorphic ...
(over 20,000 species)
Lebiinae – including Cyclosominae, Mormolycinae, Odacanthinae, Perigoninae (sometimes in Harpalinae)
Licininae – including Chlaeniinae, Oodinae (sometimes in Harpalinae)
Melaeninae
Migadopinae
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 ...
(sometimes in Harpalinae)
Panagaeinae (sometimes in Harpalinae)
Platyninae (sometimes in Harpalinae)
Pseudomorphinae
Pseudomorphinae is a subfamily of ground beetles (family
Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to ...
(sometimes in Harpalinae)
Psydrinae
Psydrinae is a subfamily of beetles in the family Carabidae.
Genera
The subfamily contains the following genera:
* ''Amblytelus
''Amblytelus'' is a genus of ground beetle including 47 species distributed through southern Australia, including ...
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 (sometimes in Harpalinae)
Trechinae
Trechinae is a subfamily in the ground beetle family, Carabidae.
Genera
The subfamily includes the following genera:
* '' Accoella'' Uéno, 1990
* '' Acheroniotes'' Lohai & Lakota, 2010
* '' Adriaphaenops'' Noesske, 1928
* '' Aepiblemus'' Bel ...
– including Bembidiinae, Patrobinae
Tribes ''incertae sedis
' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
''
* Amarotypini – Migadopinae or a distinct subfamily?
* Gehringiini – Psydrinae, Trechinae, or a distinct subfamily?
Notes
References
Further reading
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External links
Carabidae of the World
{{DEFAULTSORT:Ground Beetle