Ground beetle

Ground beetle

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Encyclopedia
Ground beetles are a large, cosmopolitan
Cosmopolitan distribution
In biogeography, a taxon is said to have a cosmopolitan distribution if its range extends across all or most of the world in appropriate habitats. For instance, the killer whale has a cosmopolitan distribution, extending over most of the world's oceans. Other examples include humans, the lichen...

 family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 of beetle
Beetle
Coleoptera is an order of insects commonly called beetles. The word "coleoptera" is from the Greek , koleos, "sheath"; and , pteron, "wing", thus "sheathed wing". Coleoptera contains more species than any other order, constituting almost 25% of all known life-forms...

s, Carabidae, with more than 40,000 species worldwide, approximately 2,000 of which are found in North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...

 and 2,700 in Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

.

Description and ecology


Although there is some variation in their body shape and coloring, most are shiny black or metallic and have ridged wing covers (elytra). The elytra are fused in some species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

, particularly large Carabinae
Carabinae
Carabinae is a subfamily of beetles in the family Carabidae, containing the following genera:* Aplothorax Waterhouse, 1841* Calosoma Weber, 1801* Carabus Linnaeus, 1758* Ceroglossus Solier, 1848* Cychropsis Boileau, 1901...

, rendering the beetles unable to fly. The genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 Mormolyce is known as violin beetle
Violin beetle
Violin beetles are ground beetles in the subfamily Lebiinae. They live between layers of bracket fungi.There are five species all contained in the genus Mormolyce.*Mormolyce castelnaudi Deyrolle, 1862 — Malaysia, Thailand...

s due to their peculiarly shaped elytra. All carabids except the quite primitive flanged bombardier beetles (Paussinae) have a groove on their foreleg tibiae
Arthropod leg
The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa , trochanter , femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus ,...

 bearing a comb of hairs. This is used for cleaning their antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....

.


Defensive secretions


Typical for the ancient beetle suborder Adephaga
Adephaga
Adephaga , with more than 40,000 recorded species in 10 families, is a suborder of highly specialized beetles and the second largest suborder of the order Coleoptera. Members of this suborder are adephagans, a term which notably include ground beetles, tiger beetles, predacious diving beetles, and...

 to which they belong, they have paired pygidial
Pygidium
The pygidium is the posterior body part or shield of crustaceans and some other arthropods, such as insects and the extinct trilobites. It contains the anus and, in females, the ovipositor...

 gland
Gland
A gland is an organ in an animal's body that synthesizes a substance for release of substances such as hormones or breast milk, often into the bloodstream or into cavities inside the body or its outer surface .- Types :...

s in the lower back of the abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...

. These are well developed in ground beetles, and produce noxious or even caustic secretions used to deter would-be predators. In some, commonly known as bombardier beetle
Bombardier beetle
Bombardier beetles are ground beetles in the tribes Brachinini, Paussini, Ozaenini, or Metriini—more than 500 species altogether—which are most notable for the defense mechanism that gives them their name: When disturbed, the beetle ejects a noxious chemical spray in a rapid burst of pulses from...

s, these secretions are mixed with volatile
Volatility (chemistry)
In chemistry and physics, volatility is the tendency of a substance to vaporize. Volatility is directly related to a substance's vapor pressure. At a given temperature, a substance with higher vapor pressure vaporizes more readily than a substance with a lower vapor pressure.The term is primarily...

 compounds and ejected by a small combustion
Combustion
Combustion or burning is the sequence of exothermic chemical reactions between a fuel and an oxidant accompanied by the production of heat and conversion of chemical species. The release of heat can result in the production of light in the form of either glowing or a flame...

, producing a loud popping sound and a cloud of hot and acrid gas which can injure small mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s like shrew
Shrew
A shrew or shrew mouse is a small molelike mammal classified in the order Soricomorpha. True shrews are also not to be confused with West Indies shrews, treeshrews, otter shrews, or elephant shrews, which belong to different families or orders.Although its external appearance is generally that of...

s, and is liable to kill invertebrate
Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...

 predators outright. To humans, getting "bombed" by a bombardier beetle is a decidedly unpleasant experience. The "bombing" ability has evolved
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

 independently twice as it seems – in the flanged bombardier beetles (Paussinae) which are among the most ancient ground beetles, as well as in the typical bombardier beetles (Brachininae
Brachininae
Brachininae is a subfamily of beetles in the family Carabidae, containing the following genera:* Aptinoderus Hubenthal, 1919* Aptinus Bonelli, 1810* Brachinulus Basilewsky, 1958* Brachinus Weber, 1801* Brachynillus Reitter, 1904...

) which are part of a more "modern" lineage. The Anthiini, meanwhile, can mechanically squirt their defensive secretions for considerable distances and are able to aim with a startling degree of accuracy; in Afrikaans
Afrikaans
Afrikaans is a West Germanic language, spoken natively in South Africa and Namibia. It is a daughter language of Dutch, originating in its 17th century dialects, collectively referred to as Cape Dutch .Afrikaans is a daughter language of Dutch; see , , , , , .Afrikaans was historically called Cape...

 they are known as ("eye-pissers"). In one of the very few known cases of a vertebrate
Vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...

 mimicking an arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...

, juvenile Heliobolus lugubris lizard
Lizard
Lizards are a widespread group of squamate reptiles, with nearly 3800 species, ranging across all continents except Antarctica as well as most oceanic island chains...

s are colored similar to the aposematic  beetles, and move in a way that makes them look surprisingly similar to the insects at a casual glance.

It is sometimes suggested that Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

 found himself on the receiving end of a bombardier beetle's defences on a collecting trip in 1828, but this is based on a misreading of his autobiography
Autobiography
An autobiography is a book about the life of a person, written by that person.-Origin of the term:...

; a bombardier beetles' "bombing" is already triggered by picking it up, and Darwin had been carrying the beetle in question in his closed hand for some time already before he ran afoul of its secretions. He discussed this incident and another such case in a letter to Leonard Jenyns as follows:
"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
River Cam
The River Cam is a tributary of the River Great Ouse in the east of England. The two rivers join to the south of Ely at Pope's Corner. The Great Ouse connects the Cam to England's canal system and to the North Sea at King's Lynn...

 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
Panagaeus cruxmajor
The crucifix ground beetle, Panagaeus cruxmajor, is a rare European ground beetle. In England it occurs in a few places only. Panagaeus bipustulatus is a commoner relative, looking very much alike except for being smaller. The crucifix ground beetle is sometimes included in P...

; 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
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...

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
Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...

 prey they can overpower. Some will run swiftly to catch their prey; tiger beetle
Tiger beetle
The tiger beetles are a large group of beetles known for their aggressive predatory habits and running speed. The fastest species of tiger beetle can run at a speed of 9 km/h , which, relative to its body length, is about 22 times the speed of former Olympic sprinter Michael Johnson, the...

s (Cicindelinae) can sustain speeds of 8 km/h (5 mph) – in relation to their body length they are among the very 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
Harpaphe haydeniana
Harpaphe haydeniana is a millipede found in the moist forests along the Pacific coast of North America, from California to British Columbia and Alaska....

, countering the hydrogen cyanide which makes these millipedes poisonous to most carnivores.

Relationship with humans


As predators of invertebrate
Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...

s, including many pests, most ground beetles are considered beneficial organisms. The caterpillar hunters (Calosoma) are famous for their habit of devouring insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

 larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...

e and pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...

e in quantity, eagerly feeding on tussock moth (Lymantriidae) caterpillars, processionary caterpillars (Thaumetopoeidae) and woolly worms (Arctiidae), which due to their urticating hair
Urticating hair
Urticating hairs, i.e. stinging hairs, are one of the primary defense mechanisms used by numerous plants, some New World tarantulas, and various lepidopteran caterpillars. Urtica is Latin for "nettle", and hairs that urticate are characteristic of this type of plant, and many other plants in...

s are avoided by most insectivore
Insectivore
An insectivore is a type of carnivore with a diet that consists chiefly of insects and similar small creatures. An alternate term is entomophage, which also refers to the human practice of eating insects....

s. Large numbers of the Forest Caterpillar Hunter (C. sycophanta), native to Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

, were shipped to New England
New England
New England is a region in the northeastern corner of the United States consisting of the six states of Maine, New Hampshire, Vermont, Massachusetts, Rhode Island, and Connecticut...

 for biological control
Biological pest control
Biological control of pests in agriculture is a method of controlling pests that relies on predation, parasitism, herbivory, or other natural mechanisms...

 of the Gypsy Moth
Gypsy moth
The gypsy moth, Lymantria dispar, is a moth in the family Lymantriidae of Eurasian origin. Originally ranging from Europe to Asia, it was introduced to North America in the late 1860s and has been expanding its range ever since...

 (Lymantria dispar) as early as 1905.

A few species are nuisance pests. Zabrus
Zabrus
Zabrus is a genus of ground beetles. They are, unusual for ground beetles, omnivores or even herbivores, and Zabrus tenebrioides can become a pest in cereal fields.-External links:*...

 is one of the few herbivorous ground beetle genera, and on rare occasions Zabrus tenebrioides for example occurs abundantly enough to cause some damage to grain
GRAIN
GRAIN is a small international non-profit organisation that works to support small farmers and social movements in their struggles for community-controlled and biodiversity-based food systems. Our support takes the form of independent research and analysis, networking at local, regional and...

 crops. Large species, usually Carabinae
Carabinae
Carabinae is a subfamily of beetles in the family Carabidae, containing the following genera:* Aplothorax Waterhouse, 1841* Calosoma Weber, 1801* Carabus Linnaeus, 1758* Ceroglossus Solier, 1848* Cychropsis Boileau, 1901...

, can become a nuisance if present in numbers, particularly during outdoor activities such as camping
Camping
Camping is an outdoor recreational activity. The participants leave urban areas, their home region, or civilization and enjoy nature while spending one or several nights outdoors, usually at a campsite. Camping may involve the use of a tent, caravan, motorhome, cabin, a primitive structure, or no...

; they will void their defensive secretions when threatened, and if they hide among provisions this can despoil food. Since ground beetles are generally reluctant or even unable to fly, it is usually easy to block their potential routes of entry mechanically or with a topical
Topical
In medicine, a topical medication is applied to body surfaces such as the skin or mucous membranes such as the vagina, anus, throat, eyes and ears.Many topical medications are epicutaneous, meaning that they are applied directly to the skin...

 insecticide
Insecticide
An insecticide is a pesticide used against insects. They include ovicides and larvicides used against the eggs and larvae of insects respectively. Insecticides are used in agriculture, medicine, industry and the household. The use of insecticides is believed to be one of the major factors behind...

.


Especially in the 19th century and to a lesser extent today, their large size and conspicuous coloration as well as the odd morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

 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 there was a veritable "beetle craze" in England
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...

. As mentioned above, Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

 was an ardent collector of beetles when he was about twenty years old, to the extent that he'd rather scour the countryside for rare specimens with William Darwin Fox
William Darwin Fox
The Reverend William Darwin Fox was an English clergyman, naturalist, and a 2nd cousin of Charles Robert Darwin.- Early life :...

, John Stevens Henslow
John Stevens Henslow
John Stevens Henslow was an English clergyman, botanist and geologist. He is best remembered as friend and mentor to his pupil Charles Darwin.- Early life :...

 and Henry Thompson
Sir Henry Thompson, 1st Baronet
Sir Henry Thompson, 1st Baronet FRCS , British surgeon and polymath, was born at Framlingham, Suffolk.-Medical career:...

 than to study theology
Theology
Theology is the systematic and rational study of religion and its influences and of the nature of religious truths, or the learned profession acquired by completing specialized training in religious studies, usually at a university or school of divinity or seminary.-Definition:Augustine of Hippo...

 as his father wanted him to do. In his autobiography
Autobiography
An autobiography is a book about the life of a person, written by that person.-Origin of the term:...

 he fondly recalled his experiences with Licinus
Licinus
Licinus is a genus of ground beetles in the Carabidae family native to the Palearctic , the Near East and North Africa. It contains the following species:* Licinus aegyptiacus Dejean, 1826...

 and Panagaeus
Panagaeus
Panagaeus is a genus of ground beetle native to the Palearctic , the Near East, and North Africa. It contains the following species:* Panagaeus abei Nakane, 1997* Panagaeus asuai Ogueta, 1966...

, 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
Adephaga
Adephaga , with more than 40,000 recorded species in 10 families, is a suborder of highly specialized beetles and the second largest suborder of the order Coleoptera. Members of this suborder are adephagans, a term which notably include ground beetles, tiger beetles, predacious diving beetles, and...

 are documented since the end of the Permian
Permian
The PermianThe term "Permian" was introduced into geology in 1841 by Sir Sir R. I. Murchison, president of the Geological Society of London, who identified typical strata in extensive Russian explorations undertaken with Edouard de Verneuil; Murchison asserted in 1841 that he named his "Permian...

, about . Ground beetles evolved
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

 in the latter Triassic
Triassic
The Triassic is a geologic period and system that extends from about 250 to 200 Mya . As the first period of the Mesozoic Era, the Triassic follows the Permian and is followed by the Jurassic. Both the start and end of the Triassic are marked by major extinction events...

, having separated from their closest relatives by . The family diversified throughout the Jurassic
Jurassic
The Jurassic is a geologic period and system that extends from about Mya to  Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...

, and the more advanced lineages, such as the Harpalinae
Harpalinae
Harpalinae is a huge subfamily of ground beetles. Among the more than 20,000 described species in this clade are the sun beetles and the violin beetles . The Harpalinae contain the most apomorphic ground beetles, displaying a wide range of forms and behaviors...

, underwent a vigorous radiation starting in the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

. The closest living relatives of the ground beetles are the false ground beetle
False ground beetle
The Trachypachidae are a family of beetles that generally resemble small ground beetles, but that are distinguished by the large coxae of their rearmost legs...

s (Trachypachidae) and the wrinkled bark beetle
Wrinkled bark beetle
Rhysodidae is a family of beetles, consisting of several hundred species in about 20 genera.These beetles are elongate, in size ranging from 5–8 mm, and color ranging from a reddish brown to black. Both the thorax and the elytra are deeply grooved lengthwise, thus giving these beetles their...

s (Rhysodidae). They are sometimes even included in the Carabidae as subfamilies or as tribes incertae sedis
Incertae sedis
, is a term used to define a taxonomic group where its broader relationships are unknown or undefined. Uncertainty at specific taxonomic levels is attributed by , , and similar terms.-Examples:*The fossil plant Paradinandra suecica could not be assigned to any...

, but more preferably they are united with the ground beetles in the superfamily
Taxonomic rank
In biological classification, rank is the level in a taxonomic hierarchy. Examples of taxonomic ranks are species, genus, family, and class. Each rank subsumes under it a number of less general categories...

 Caraboidea.

Much research has been done on elucidating the phylogeny of the ground beetles and adjusting systematics
Systematics
Biological systematics is the study of the diversification of terrestrial life, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees...

 and taxonomy
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...

 accordingly. While there is no completely firm consensus, a few points are generally accepted: As it seems, the ground beetles consist of a number of more basal lineages and the extremely diverse Harpalinae
Harpalinae
Harpalinae is a huge subfamily of ground beetles. Among the more than 20,000 described species in this clade are the sun beetles and the violin beetles . The Harpalinae contain the most apomorphic ground beetles, displaying a wide range of forms and behaviors...

 which contain over half the described species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 and into which several formerly independent families had to be subsumed.

Subfamilies and selected genera


The taxonomy
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...

 used here is 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
Tree of Life Web Project
The Tree of Life Web Project is an ongoing 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....

 makes little use of subfamilies, listing most tribe
Tribe (biology)
In biology, a tribe is a taxonomic rank between family and genus. It is sometimes subdivided into subtribes.Some examples include the tribes: Canini, Acalypheae, Hominini, Bombini, and Antidesmeae.-See also:* Biological classification* Rank...

s as incertae sedis
Incertae sedis
, is a term used to define a taxonomic group where its broader relationships are unknown or undefined. Uncertainty at specific taxonomic levels is attributed by , , and similar terms.-Examples:*The fossil plant Paradinandra suecica could not be assigned to any...

 as to subfamily. Fauna Europaea on the other hand splits rather than lumps
Lumpers and splitters
Lumping and splitting refers to a well-known problem in any discipline which has to place individual examples into rigorously defined categories. The lumper/splitter problem occurs when there is the need to create classifications and assign examples to them, for example schools of literature,...

 the Harpalinae, restricting them to what in the system uses here is the tribe Harpalini.

All the approaches mentioned above are legitimate as they agree with the phylogeny as far as it has been resolved. The inclusive Harpalinae presented here are used for two reasons, one scientific and one practical – first, the majority of authors presently uses this system, following the Catalogue of Palaearctic Coleoptera. Second, the MediaWiki
MediaWiki
MediaWiki is a popular free web-based wiki software application. Developed by the Wikimedia Foundation, it is used to run all of its projects, including Wikipedia, Wiktionary and Wikinews. Numerous other wikis around the world also use it to power their websites...

 markup
Markup language
A markup language is a modern system for annotating a text in a way that is syntactically distinguishable from that text. The idea and terminology evolved from the "marking up" of manuscripts, i.e. the revision instructions by editors, traditionally written with a blue pencil on authors' manuscripts...

 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
Carabinae is a subfamily of beetles in the family Carabidae, containing the following genera:* Aplothorax Waterhouse, 1841* Calosoma Weber, 1801* Carabus Linnaeus, 1758* Ceroglossus Solier, 1848* Cychropsis Boileau, 1901...

 Latreille, 1802 – including Agoninae and Callistinae
  • Altagonum
  • Aplothorax
    Aplothorax
    Aplothorax burchelli is a species of beetle in the family Carabidae, the only species in the genus Aplothorax....

     (monotypic genus)
  • Calosoma
    Calosoma
    Calosoma is a genus of large ground beetles that occur primarily throughout the Northern Hemisphere, and are referred to as caterpillar hunters or searchers. Many of the 167 species are largely or entirely black, but some have bright metallic coloration...

     - including Callisthenes
  • Carabus
    Carabus
    Carabus is a genus of beetle in family Carabidae.-Species:* Carabus aba Kalab, 2002* Carabus abbreviatus Brulle, 1835* Carabus absonus Cavazzuti & Rapuzzi, 2005* Carabus achilleanus Cavazzuti, 2002...

  • Cychrus
    Cychrus
    Cychrus is a genus of beetles in the family Carabidae, containing the following species:* Cychrus aeneus Fisher-Waldheim, 1824* Cychrus anatolicus Motschulsky, 1865* Cychrus andrei Cavazzuti, 2001* Cychrus angulicollis Sella, 1874...

  • Homothes
  • Fortagonum
  • Laemostenus
    Laemostenus
    Laemostenus is a genus of ground beetles present on all continents on Earth, except Antarctica.-Selected subgenera and species:*Laemostenus **Laemostenus acutangulus**Laemostenus angustatus**Laemostenus carinatus...

  • Notagonum


Cicindelinae – tiger beetles (roughly 2,100 species; sometimes included in Carabidae)

Cicindinae

Elaphrinae
Elaphrinae
Elaphrinae is a subfamily of beetles in the family Carabidae, containing the following genera:* Blethisa Bonelli, 1810* Diacheila Motschulsky, 1844* Elaphrus Fabricius, 1775...

 Latreille, 1802
  • Blethisa
    Blethisa
    Blethisa is a genus of ground beetle native to the Palearctic. It contains the following species:* Blethisa catenaria Brown, 1944* Blethisa eschscholtzii Zoubkoff, 1829* Blethisa inexpectata Goulet & Smetana, 1983...

  • Diacheila
    Diacheila
    Diacheila is a genus of beetles in the family Carabidae, containing the following species:* Diacheila arctica Gyllenhal, 1810* Diacheila fausti Heyden, 1887* Diacheila polita Faldermann, 1835...

     Motschulsky, 1844
  • Elaphrus
    Elaphrus
    Elaphrus is a genus of ground beetle native to the Palearctic, the Nearctic, the Near East and Northern Africa. It contains the following species:* Elaphrus americanus Dejean, 1831 * Elaphrus angulonotus Shi & Liang, 2008...



Hiletinae
Hiletinae
Hiletinae is a subfamily of beetles in the family Carabidae. It contains 21 species in the following two genera:* Genus Eucamaragnathus Jeannel, 1938** Eucamaragnathus amapa Erwin & Stork, 1985** Eucamaragnathus angulicollis...



Loricerinae
Loricerinae
Loricerinae is a subfamily of beetles in the family Carabidae. It contains the single genus Loricera with the following species:* Loricera aptera Ball & Erwin, 1969 * Loricera balli Sciaky & Facchini, 1999...

 Bonelli, 1810
  • Loricera
    Loricera
    Loricera is a genus of ground beetle native to the Palearctic and the Nearctic.Species include:*Loricera pilicornis-External links:*...



Migadopinae
Migadopinae
Migadopinae is a subfamily of beetles in the family Carabidae, containing the following genera:* Amarotypus Bates, 1872 * Antarctonomus Chaudoir, 1861 * Aquilex Moret, 1990 * Calathosoma Jeannel, 1938 * Calyptogonia Sloane, 1920...



Nebriinae
Nebriinae
Nebriinae is a subfamily of beetles in the family Carabidae, containing the following genera:* Archaeocindis Kavanaugh & Erwin, 1991 * Archastes Jedlicka, 1935* Archileistobrius Shilenkov & Kryzhanovskij, 1983 * Cicindis Bruch, 1908...

 (includes Notiophilinae, often included in Carabinae)
  • Leistus
    Leistus
    Leistus is a genus of ground beetle native to the Palearctic and the Nearctic. It contains the following species:* Leistus acutangulus Perrault, 1979* Leistus alaiensis Kabak, 1995* Leistus andrewesi Perrault, 1985...

  • Nebria
    Nebria
    Nebria is a genus of ground beetle native to the Palearctic, the Near East and North Africa. It contains the following species:* Nebria aborana Andrewes, 1925* Nebria acuta Lindroth, 1961 * Nebria adjarica Shilenkov, 1983...

  • Notiophilus
    Notiophilus
    Notiophilus is a genus of ground beetle native to the Palearctic, the Nearctic, the Near East and North Africa. It contains the following species:* Notiophilus aeneus * Notiophilus aestuans Dejean, 1826...

  • Pelophila
    Pelophila
    Pelophila is a genus of beetles in the family Carabidae, containing the following species:* Pelophila borealis * Pelophila rudis LeConte, 1863...

     Dejean, 1821


Nototylinae
Nototylinae
Nototylinae is a subfamily of beetles in the family Carabidae. It contains the single genus Nototylus with the single species Nototylus fryi....



Omophroninae
Omophroninae
Omophroninae is a subfamily of beetles in the family Carabidae. It contains the single genus Omophron with the following species:* Omophron aequale A...

 Bonelli, 1810 – round sand beetles
  • Omophron
    Omophron
    Omophron is a genus of ground beetle native to the Palearctic, the Near East and North Africa.Species include:*Omophron limbatum-External links:*...



Paussinae – ant nest beetles, flanged bombardier beetles

Promecognathinae

Scaritinae Bonelli, 1810 – pedunculate ground beetles
  • Clivina
    Clivina
    Clivina is a genus of ground beetle native to the Palearctic, the Nearctic, the Near East and North Africa.-External links:*...

  • Dyschirius Bonelli, 1810
  • Scarites
    Scarites
    Scarites is a genus of ground beetle native to the Palearctic, the Near East, North America and North Africa. These beetles share physical characteristics of the more tropical stag beetles, but are not closely related. Scarites can often be found under loose rocks and boards. If touched, they often...



Siagoninae Bonelli, 1810

Carabidae Conjunctae




Amblytelinae Sloane, 1898
  • Amblytelus
    Amblytelus
    Amblytelus is a genus of ground beetle including 47 species distributed through southern Australia, including the Southwest and along the east coast up to North Queensland.-External links:*...



Apotominae
Apotominae
Apotominae is a subfamily of beetles in the family Carabidae. It contains the single genus Apotomus with the following species:* Apotomus alluaudi Jeannel, 1946* Apotomus angusticollis J...

  • Apotomus Illiger, 1807


Brachininae
Brachininae
Brachininae is a subfamily of beetles in the family Carabidae, containing the following genera:* Aptinoderus Hubenthal, 1919* Aptinus Bonelli, 1810* Brachinulus Basilewsky, 1958* Brachinus Weber, 1801* Brachynillus Reitter, 1904...

 Bonelli, 1810 – typical bombardier beetles
  • Aptinus
    Aptinus
    Aptinus is a genus of ground beetle native to Europe and the Near East. It contains the following species:* Aptinus acutangulus Chaudoir, 1876* Aptinus alpinus Dejean, 1829* Aptinus bombarda...

     (tentatively placed here)
  • Brachinus
    Brachinus
    Brachinus is a genus of ground beetle native to the Palearctic, the Near East and North Africa. It contains the following species:* Brachinus aabaaba Erwin, 1970* Brachinus abbreviatus...

  • Mastax
    Mastax
    Mastax is a genus of beetles in the family Carabidae, containing the following species:* Mastax albonotata Peringuey, 1885* Mastax alternans Basilewsky, 1959* Mastax annulata Andrewes, 1924* Mastax brittoni Quentin, 1952...

     Fischer von Waldheim, 1828


Broscinae
Broscinae
Broscinae is a subfamily of beetles in the family Carabidae, containing the following genera:* Acallistus Sharp, 1886* Adoleta Castelnau, 1867* Anheretus Putzeys, 1868* Axonya Andrewes, 1923* Barypus Dejean, 1828...

 Hope, 1838

  • Acallistus
    Acallistus
    Acallistus is a genus of beetles in the family Carabidae, containing the following species:* Acallistus cuprescens * Acallistus longus * Acallistus plebius...

     Sharp, 1886
  • Adotela Laporte de Castelnau, 1867
  • Axonya
    Axonya
    Axonya championi is a species of beetle in the family Carabidae, the only species in the genus Axonya....

     Andrewes, 1923
  • Baripus Dejean, 1828
  • Bountya
    Bountya
    Bountya insularis is a species of beetle in the family Carabidae, the only species in the genus Bountya....

     Townsend, 1971
  • Brithysternum Macleay, 1873
  • Broscodera
    Broscodera
    Broscodera is a genus of beetles in the family Carabidae, containing the following species:* Broscodera insignis Mannerheim, 1852* Broscodera dreuxi Deuve, 1990* Broscodera holzschuhi Wrase, 1995...

     Lindroth, 1961
  • Broscodes
    Broscodes
    Broscodes karumicus is a species of beetle in the family Carabidae, the only species in the genus Broscodes....

     Bolivar, 1914
  • Broscosoma
    Broscosoma
    Broscosoma is a genus of beetles in the family Carabidae, containing the following species:* Broscosoma baldense Rosenhauer, 1846* Broscosoma businskae Dvorak, 1998* Broscosoma convexum Deuve, 1983* Broscosoma deuvei Lassalle, 1982...

     Rosenhauer, 1846
  • Broscus
    Broscus
    Broscus is a genus of ground beetle native to the Palearctic, the Nearctic, the Near East and North Africa. It contains the following species:*Broscus aberti Jedlicka, 1965*Broscus asiaticus Ballion, 1871*Broscus augustulus Semenov, 1891...

  • Brullea
    Brullea
    Brullea antarctica is a species of beetle in the family Carabidae, the only species in the genus Brullea....

     Laporte de Castelnau, 1868
  • Cascellius
    Cascellius
    Cascellius is a genus of beetles in the family Carabidae, containing the following species:* Cascellius gravesii Curtis, 1839* Cascellius septentrionalis Roig-Junent, 1995...

     Curtis, 1839

  • Cerotalis
    Cerotalis
    Cerotalis is a genus of beetles in the family Carabidae, containing the following species:* Cerotalis amabilis Sloane, 1890* Cerotalis brachypleura Sloane, 1898* Cerotalis longipes Sloane, 1898* Cerotalis majuscula...

     Laporte de Castelnau, 1868
  • Chaetobroscus
    Chaetobroscus
    Chaetobroscus is a genus of beetles in the family Carabidae, containing the following species:* Chaetobroscus anomalus Chaudoir, 1878* Chaetobroscus kezukai Dostal, 1984* Chaetobroscus bhutanensis Morvan, 1980...

     Semenov, 1900
  • Chylnus
    Chylnus
    Chylnus is a genus of beetles in the family Carabidae, containing the following species:* Chylnus ater * Chylnus concolor * Chylnus montanum * Chylnus substriatum...

     Sloane, 1920
  • Craspedonotus
    Craspedonotus
    Craspedonotus is a genus of beetles in the family Carabidae, containing the following species:* Craspedonotus himalayanus Semenov, 1910* Craspedonotus margellanicus Kraatz, 1884* Craspedonotus tibialis Schaum, 1863...

     Schaum, 1863
  • Creobius
    Creobius
    Creobius eudouxii is a species of beetle in the family Carabidae, the only species in the genus Creobius....

     Guérin-Méneville, 1838
  • Diglymma
    Diglymma
    Diglymma is a genus of beetles in the family Carabidae, containing the following species:* Diglymma castigatum Broun, 1909* Diglymma clivinoides * Diglymma marginale Broun, 1914* Diglymma obtusum...

     Sharp, 1886
  • Eobroscus
    Eobroscus
    Eobroscus is a genus of beetles in the family Carabidae, containing the following species:* Eobroscus bhutanensis Morvan, 1982* Eobroscus lutshniki Roubal, 1928* Eobroscus masumotoi Morita, 1990* Eobroscus uenoi Morita, 1995...

     Kruizhanovskií, 1951
  • Eurylychnus
    Eurylychnus
    Eurylychnus is a genus of beetles in the family Carabidae, containing the following species:* Eurylychnus blagravii * Eurylychnus cylindricus Sloane, 1916* Eurylychnus dyschirioides...

     Bates, 1891
  • Gnathoxys
    Gnathoxys
    Gnathoxys is a genus of beetles in the family Carabidae, containing the following species:* Gnathoxys barbatus MacLeay, 1864* Gnathoxys cicatricosus Reiche, 1842* Gnathoxys crassipes Sloane, 1898* Gnathoxys foveatus MacLeay, 1863...

     Westwood, 1842
  • Mecodema
    Mecodema
    Mecodema is a genus of ground beetle in the family Carabidae.-Species:* Mecodema allani Fairburn, 1945* Mecodema alternans Laporte de Castelnau, 1867* Mecodema angustulum Broun, 1914* Mecodema aoteanoho Seldon & Leschnen, 2011...

  • Metaglymma
    Metaglymma
    Metaglymma is a genus of beetles in the family Carabidae, containing the following species:* Metaglymma aberrans Putzeys, 1868* Metaglymma moniliforme Bates, 1867* Metaglymma tibiale...

     Bates, 1867

  • Microbarypus
    Microbarypus
    Microbarypus silvicola is a species of beetle in the family Carabidae, the only species in the genus Microbarypus....

     Roig-Juñent, 2000
  • Miscodera
    Miscodera
    Miscodera arctica is a species of beetle in the family Carabidae, the only species in the genus Miscodera....

     Eschscholtz, 1830
  • Nothobroscus
    Nothobroscus
    Nothobroscus chilensis is a species of beetle in the family Carabidae, the only species in the genus Nothobroscus....

     Roig-Juñent & Ball, 1995
  • Nothocascellius
    Nothocascellius
    Nothocascellius is a genus of beetles in the family Carabidae, containing the following species:* Nothocascellius aeneoniger * Nothocascellius hyadesii...

     Roig-Juñent, 1995
  • Oregus
    Oregus
    Oregus is a genus of beetles in the family Carabidae, containing the following species:* Oregus aereus * Oregus inaequalis...

     Putzeys, 1868
  • Parroa Laporte de Castelnau, 1868
  • Percolestus
    Percolestus
    Percolestus blackburni is a species of beetle in the family Carabidae, the only species in the genus Percolestus....

     Sloane, 1892
  • Percosoma
    Percosoma
    Percosoma is a genus of beetles in the family Carabidae, containing the following species:* Percosoma asymetricum Fauvel, 1903* Percosoma carenoides * Percosoma sulcipenne Bates, 1878...

     Schaum
    Hermann Rudolph Schaum
    Hermann Rudolph Schaum was a professor in Berlin and an entomologist. He specialised in Coleoptera....

    , 1858
  • Promecoderus Dejean, 1829
  • Rawlinsius
    Rawlinsius
    Rawlinsius papillatus is a species of beetle in the family Carabidae, the only species in the genus Rawlinsius....

     Davidson & Ball, 1998
  • Zacotus
    Zacotus
    Zacotus matthewsii is a species of beetle in the family Carabidae, the only species in the genus Zacotus....

     Leconte, 1869




Harpalinae
Harpalinae
Harpalinae is a huge subfamily of ground beetles. Among the more than 20,000 described species in this clade are the sun beetles and the violin beetles . The Harpalinae contain the most apomorphic ground beetles, displaying a wide range of forms and behaviors...

 – including Chlaeniinae, Cyclosominae, Dryptinae, Lebiinae, Licininae, Mormolycinae, Odacanthinae, Oodinae, Panagaeinae, Perigoninae, Platyninae, Pseudomorphinae, Pterostichinae, Zabrinae (over 20,000 species)

Melaeninae
Melaeninae
Melaeninae is a subfamily of beetles in the family Carabidae. It contains 23 species in 2 genera:* Genus Cymbionotum Baudi di Selva, 1864** Cymbionotum basale Dejean, 1831** Cymbionotum candidum Andrewes, 1935...



Psydrinae
  • Mecyclothorax


Trechinae Bonelli, 1810 – including Bembidiinae, Patrobinae

  • Aepus Samouelle, 1819
  • Amerizus de Chaudoir 1868 – including Gnatholymnaeum
  • Anillinus Casey, 1918
  • Anophthalmus
    Anophthalmus
    Anophthalmus is a genus of ground beetle endemic to Europe.-External links:*...

  • Asaphidion
    Asaphidion
    Asaphidion is a Holarctic genus of ground beetle native to Europe, the Near East, North Africa, and North America.-External links:**...

  • Bembidion
    Bembidion
    Bembidion is the largest genus of beetles in the family Carabidae by number of species. There have been many attempts to divide it into smaller genera, most notably by René Jeannel in 1941 and by G.G. Perrault in 1981, but none of them has been generally accepted. All species are small and move...

  • Blemus
    Blemus
    Blemus is a genus of ground beetle native to the Palearctic and the Nearctic.Species include:* Blemus alexandrovi * Blemus discus -External links:*...

     – including Lasiotrechus
  • Cardiaderus Dejean, 1828
  • Cillenus Leach, 1819
  • Deltomerus Motschulsky, 1850
  • Duvaliopsis Jeannel, 1928 (tentatively placed here)
  • Duvalius Delarouzée, 1859
  • Lymnastis Motschulsky
    Victor Ivanovitsch Motschulsky
    Victor Ivanovitsch Motschulsky was a Russian entomologist mainly interested in beetles....

    , 1862
  • Miscodera
    Miscodera
    Miscodera arctica is a species of beetle in the family Carabidae, the only species in the genus Miscodera....

     Eschscholtz, 1830
  • Ocys Stephens, 1828

  • Patrobus
    Patrobus
    Patrobus is a genus of ground beetle native to the Palearctic and the Nearctic.-External links:*...

  • Perileptus Schaum, 1860
  • Pogonus Dejean, 1821
  • Porotachys Netolitzky, 1914 (tentatively placed here)
  • Pseudanophthalmus
    Pseudanophthalmus
    Pseudanophthalmus is a genus of cave beetle. Over 200 species have been described in the caves of ten states of the eastern United States.-Taxonomy:The genus was first named and described by René Jeannel in 1920...

  • Pseudaphaenops Winkler, 1912
  • Serranillus Barr, 1996
  • Tachyra Motschulsky
    Victor Ivanovitsch Motschulsky
    Victor Ivanovitsch Motschulsky was a Russian entomologist mainly interested in beetles....

    , 1862
  • Tachys Dejean, 1821
  • Tachyta Kirby, 1837
  • Thalassophilus Wollaston, 1854
  • Trechoblemus Ganglbauer, 1891
  • Trechosia Jeannel, 1926
  • Trechus
    Trechus
    Trechus is a genus of ground beetle native to the Palearctic and the Near East. The name is derived from the greek word trécho, meaning "I run" .-Externhal links:*...

  • Winklerites Jeannel, 1937




Tribes incertae sedis
Incertae sedis
, is a term used to define a taxonomic group where its broader relationships are unknown or undefined. Uncertainty at specific taxonomic levels is attributed by , , and similar terms.-Examples:*The fossil plant Paradinandra suecica could not be assigned to any...

  • Amarotypini
    Amarotypini
    -Introduction:Amarotypines are a small tribe of ground beetles , recognised as a distinct tribe since 1985. The best known species is Amarotypus edwardsii of New Zealand, widely distributed throughout the three main islands, and arboreal in habits.-Biodiversity and distribution:* Genus Amarotypus:...

     – Harpalinae?
  • Gehringiini – Trechinae or a distinct subfamily
  • Metiini – Harpalinae?

External links