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 enti ...
. As of 2015, it is one of the 10 most species-rich animal families. They belong to the
Adephaga
The Adephaga (from Greek ἀδηφάγος, ''adephagos'', "gluttonous") are a suborder of beetles, and with more than 40,000 recorded species in 10 families, the second-largest of the four beetle suborders. Members of this suborder are collecti ...
. 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 altern ...
). The elytra are fused in some
species
In biology, a species is the basic unit of Taxonomy (biology), 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 ...
, 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'' Soli ...
, rendering the beetles unable to fly. The
species
In biology, a species is the basic unit of Taxonomy (biology), 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 ...
''
Mormolyce phyllodes
''Mormolyce phyllodes'', commonly known as the violin beetle, is a species of ground beetles in the subfamily Lebiinae.
Subspecies
The species may be divided into the following subspecies:Lorenz, W Nomina Carabideum, Online Database"/ref>
* ...
'' is known as violin beetle due to their peculiarly shaped elytra. All carabids except the quite primitive
flanged bombardier beetle
Ant nest beetles (subfamily Paussinae) or paussines, some members of which are known also as flanged bombardier beetles, are a large subfamily within the ground beetles (Carabidae).The tribes Metriini, Ozaenini, Paussini and Protopaussini are in ...
s (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
Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
Adephaga
The Adephaga (from Greek ἀδηφάγος, ''adephagos'', "gluttonous") are a suborder of beetles, and with more than 40,000 recorded species in 10 families, the second-largest of the four beetle suborders. Members of this suborder are collecti ...
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 compo ...
gland
In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream ( endocrine gland) or into cavities inside the body or its outer surface ( exocrine gland).
Structure
...
s 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 tors ...
. 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 ''caust ...
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 ...
s. In some, commonly known as
bombardier beetle
Bombardier beetles are ground beetles (Carabidae) 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, they eje ...
s, these secretions are mixed with
volatile
Volatility or volatile may refer to:
Chemistry
* Volatility (chemistry), a measuring tendency of a substance or liquid to vaporize easily
* Relative volatility, a measure of vapor pressures of the components in a liquid mixture
* Volatiles, a gr ...
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. Combust ...
, 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 diffe ...
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 te ...
independently twice, as it seems, in the
flanged bombardier beetle
Ant nest beetles (subfamily Paussinae) or paussines, some members of which are known also as flanged bombardier beetles, are a large subfamily within the ground beetles (Carabidae).The tribes Metriini, Ozaenini, Paussini and Protopaussini are in ...
s (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.
Taxonomy
The subfamily contains two tribes and 14 genera.
* Tribe Brachinini Bonelli, 1810
** ''Aptinoderus'' Hubenthal, 1919
** '' Aptinus'' Bonelli, 1810
** ''Brachinulus'' Basil ...
), 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 g ...
, they are known as ' ("eye-pissers"). In one of the very few known cases of a
vertebrate
Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
mimicking
In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry ...
an
arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
, juvenile ''
Heliobolus lugubris
''Heliobolus lugubris'', also known commonly as the bushveld lizard, mourning racerunner, or the black and yellow sand lizard, is a species of lizard in the family Lacertidae. The species is found in Southern Africa: southern Angola, Namibia, Bot ...
''
lizard
Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia al ...
s 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 natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
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
''Cychrus caraboides'' is a species of ground beetle.
It is found in Belarus, Benelux, Great Britain including the Isle of Man, Estonia, Finland, mainland France, Hungary, the Republic of Ireland, Kaliningrad, Latvia, Moldova, Northern Ireland, ...
'' 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
A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other ...
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
Nocturnality is an ethology, animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite.
Nocturnal creatures generally have ...
, 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
''Promecognathus laevissimus'' is a species of ground beetle 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 A ...
'' are specialised predators of the cyanide millipede ''
Harpaphe haydeniana
''Harpaphe haydeniana,'' commonly known as the yellow-spotted millipede, almond-scented millipede or cyanide millipede, is a species of polydesmidan ("flat-backed") millipede found in the moist forests along the Pacific coast of North America, fr ...
'', countering the
hydrogen cyanide
Hydrogen cyanide, sometimes called prussic acid, is a chemical compound with the formula HCN and structure . It is a colorless, extremely poisonous, and flammable liquid that boils slightly above room temperature, at . HCN is produced on a ...
that makes these millipedes poisonous to most carnivores.
Relationship with humans
As predators of invertebrates, including many
pest
Pest or The Pest may refer to:
Science and medicine
* Pest (organism), an animal or plant deemed to be detrimental to humans or human concerns
** Weed, a plant considered undesirable
* Infectious disease, an illness resulting from an infection
** ...
s, 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 c ...
(Lymantriidae) caterpillars,
processionary caterpillars (Thaumetopoeidae) and
woolly worm
The Arctiinae (formerly called the family Arctiidae) are a large and diverse subfamily of moths with around 11,000 species found all over the world, including 6,000 neotropical species.Scoble, MJ. (1995). ''The Lepidoptera: Form, Function and D ...
s (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 gen ...
s, are avoided by most insectivores. Large numbers of the
forest caterpillar hunter
''Calosoma sycophanta'', the forest caterpillar hunter, is a species of ground beetle belonging to the family Carabidae.
Subspecies and varietas
* ''Calosoma sycophanta var. severum'' Chaudoir, 1850
* ''Calosoma sycophanta var. nigrocyaneum' ...
(''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 enti ...
, 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 to the west and by the Canadian province ...
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 als ...
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'' be ...
(''Lymantria dispar'') as early as 1905.
A few species are nuisance pests. ''
Zabrus
''Zabrus'' is a genus of ground beetles. They are, unusually for ground beetles, omnivores or even herbivores, and '' Zabrus tenebrioides'' can become a pest in cereal fields.
Subgenera
The following are subgenera of ''Zabrus'':
* ''Aulacozabru ...
'' 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 mouthpart ...
ground beetle genera, and on rare occasions ''
Zabrus tenebrioides
''Zabrus tenebrioides'' is a species of black coloured ground beetle in the Pterostichinae subfamily that can be found everywhere in Europe and the Near East.
Distributed in the steppe and forest-steppe to the southern border of Polesia. By its ...
'', for example, occurs abundantly enough to cause some damage to
grain
A grain is a small, hard, dry fruit ( caryopsis) – with or without an attached hull layer – harvested for human or animal consumption. A grain crop is a grain-producing plant. The two main types of commercial grain crops are cereals and legu ...
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'' Soli ...
, 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 natu ...
; 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.

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
Lebiini is a tribe of ground beetles in the family Carabidae. There are more than 250 genera and 4,800 described species in Lebiini.
Genera
These 251 genera belong to the tribe Lebiini:
;Subtribe Agrina Kirby, 1837
: '' Abaditicus'' Ball & Hil ...
), 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 solic ...
, 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 th ...
as his father wanted him to do. In his autobiography, he fondly recalled his experiences with ''
Licinus
''Licinus'' is a genus of ground beetles in the family Carabidae native to the Palearctic (including Europe), the Near East and North Africa. It contains the following species:
* '' Licinus aegyptiacus'' Dejean, 1826
* '' Licinus aequatus'' ...
'' and ''
Panagaeus
''Panagaeus'' is a genus of ground beetle native to the Holarctic (including Europe), the Near East, and North Africa. It contains the following species:
* ''Panagaeus abei'' Nakane, 1997
* ''Panagaeus asuai'' Ogueta, 1966
* ''Panagaeus bipus ...
'', 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
The Adephaga (from Greek ἀδηφάγος, ''adephagos'', "gluttonous") are a suborder of beetles, and with more than 40,000 recorded species in 10 families, the second-largest of the four beetle suborders. Members of this suborder are collecti ...
are documented since the end of the
Permian
The Permian ( ) is a geologic period and 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 period of the Pale ...
, 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 te ...
in the latter
Triassic
The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
, having separated from their closest relatives by 200 Mya. The family diversified throughout the
Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), 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 J ...
, 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 apomorphi ...
, 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
SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
Caraboidea, or
Geadephaga
The Adephaga (from Greek language, Greek ἀδηφάγος, ''adephagos'', "gluttonous") are a suborder of beetles, and with more than 40,000 recorded species in 10 families, the second-largest of the four beetle suborders. Members of this subor ...
.
[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
Biological systematics is the study of the diversification of living forms, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees (synonyms: cladograms, phylogenetic t ...
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
Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''.
Science
* Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure
* Basal (medicine), a minimal level that is nec ...
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 confl ...
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 European land and fresh-water animals. It serves as a standard taxonomic source for animal taxonomy within the Pan-European Species directories Inf ...
'', 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 Amblyst ...
.
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'' Bonelli, 1813
** ''Enceladus gigas'' Bonelli, 1813
* Genus ''Luperca'' Castelnau, 1840
** ''Luperca goryi'' (Guerin-Meneville ...
.
[
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 software, free and open-source wiki software. It is used on Wikipedia and almost all other Wikimedia movement, Wikimedia Website, websites, including Wiktionary, Wikimedia Commons and Wikidata; these sit ...
markup
Markup or mark-up can refer to:
* Markup language, a standardized set of notations used to annotate a plain-text document's content to give information regarding the structure of the text or instructions for how it is to be displayed
** Lightweigh ...
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'' Soli ...
Latreille, 1802 – including Agoninae and Callistinae
* ''Altagonum
''Altagonum'' is a genus of beetles in the family Carabidae, containing the following species:
* '' Altagonum avium'' Darlington, 1971
* '' Altagonum caducum'' Darlington, 1952
* '' Altagonum cheesmani'' Darlington, 1952
* '' Altagonum cracen ...
''
* ''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 ...
''
* ''Laemostenus
''Laemostenus'' is a genus of ground beetles present on all continents on Earth, except Antarctica.
There are nearly 200 species in the genus, divided into several subgenera. Beetles of the genus are about 8 to 28 millimeters long. Many are dark ...
''
* ''Notagonum
''Notagonum'' is a genus of beetles in the family Carabidae, containing the following species:
* ''Notagonum addendum'' Darlington, 1952
* ''Notagonum aitape'' Darlington, 1952
* ''Notagonum albertisi'' (Maindron, 1906)
* ''Notagonum altum'' Da ...
''
Cicindinae
Elaphrinae
Elaphrinae is a subfamily of beetles in the family Carabidae, containing the following genera:
* ''Blethisa
''Blethisa'' is a genus of ground beetle native to the Palearctic. It contains the following species:
* ''Blethisa catenaria'' Brown, ...
Latreille, 1802
* ''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 hudsonica'' Casey, 1924
* ''Blethisa inexpectata' ...
''
* ''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
References
Elaphrinae ...
'' 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. It contains the following species:
* '' Elaphrus americanus'' Dejean, 1831
* ''Elaphrus angulonotus'' Shi & Liang, 2008
* '' E ...
''
Hiletinae
Hiletinae is a subfamily of beetles in the family Carabidae, containing 21 species in two genera. All of the species in the genus ''Hiletus'', as well as 6 species in '' Eucamaragnathus'', live in Africa. The other species in ''Eucamaragnathus'' l ...
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
* '' Loricera barbarae'' ...
Bonelli, 1810
* ''Loricera
''Loricera'' is a genus of ground beetle native to the Palearctic and the Nearctic.
Species
''Loricera'' contains the following species:
*''Loricera aptera'' Ball & Erwin, 1969
*''Loricera balli'' Sciaky & Facchini, 1999
*''Loricera barbarae' ...
''
Migadopinae
The Migadopinae are 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 ...
Nebriinae
Nebriinae 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 Europe ...
(includes Notiophilinae, often included in Carabinae)
* ''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
* '' L ...
''
* ''Nebria
''Nebria'' is a genus of ground beetles native to the Palearctic, the Near East and North Africa
North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope ...
''
* ''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
*'' Notiop ...
''
* ''Pelophila
''Pelophila'' is a genus of beetles in the ground beetle
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 Europe. A ...
'' Dejean, 1821
* ''Nippononebria
''Nippononebria'' is a genus of ground beetles in the family Carabidae. There are about eight described species in ''Nippononebria''.
Species
These eight species belong to the genus ''Nippononebria'':
* ''Nippononebria altisierrae'' (Kavanaugh, ...
''
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 Eur ...
Omophroninae Bonelli, 1810 – round sand beetles
* ''Omophron
file:Omophron nitidum P1440156a.jpg, ''Omophron nitidum''
''Omophron'' is a genus of ground beetle (Carabidae), the only extant genus in the subfamily Omophroninae. It is mostly distributed in the Northern Hemisphere with the southern border runn ...
''
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 ...
– wrinkled bark beetles
Scaritinae
Scaritinae is a subfamily of beetles in the family Carabidae, containing the following genera:
* ''Acanthoscelis'' Dejean, 1825
* '' Afroclivina'' Kult, 1959
* '' Afrosyleter'' Basilewsky, 1959
* '' Akephorus'' LeConte, 1851
* '' Alpiodytes'' Jea ...
Bonelli, 1810 – pedunculate ground beetles
* ''Clivina
''Clivina'' is a genus of ground beetle native to the Palearctic, the Nearctic, the Near East
The ''Near East''; he, המזרח הקרוב; arc, ܕܢܚܐ ܩܪܒ; fa, خاور نزدیک, Xāvar-e nazdik; tr, Yakın Doğu is a geographic ...
''
* ''Dyschirius
''Dyschirius'' is a genus of beetles in the family Carabidae:
Species
This is a list of the species in ''Dyschirius'':
* '' Dyschirius abbreviatus'' Putzeys, 1846
* '' Dyschirius abditus'' Fedorenko, 1993
* '' Dyschirius addisabeba'' ( Bulir ...
'' Bonelli, 1810
* ''Scarites
''Scarites'' is a genus of ground beetle native to the Palearctic, the Near East, North America and North Africa. There are at least 190 described species in ''Scarites''.
These beetles share physical characteristics of the more tropical st ...
''
Siagoninae
Siagoninae is a subfamily of beetles in the family Carabidae. It contains 83 species in 3 genera:
* Genus ''Enceladus'' Bonelli, 1813
** ''Enceladus gigas'' Bonelli, 1813
* Genus ''Luperca'' Castelnau, 1840
** ''Luperca goryi'' (Guerin-Meneville ...
Bonelli, 1810
Carabidae Conjunctae
Amblytelinae Sloane, 1898
* ''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. It contains the following species:
* ''Amblytelus balli'' Baehr, 200 ...
''
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. Muller, 1943
* ''Apotomus annulaticornis'' ...
Brachininae
Brachininae is a subfamily of beetles in the family Carabidae.
Taxonomy
The subfamily contains two tribes and 14 genera.
* Tribe Brachinini Bonelli, 1810
** ''Aptinoderus'' Hubenthal, 1919
** '' Aptinus'' Bonelli, 1810
** ''Brachinulus'' Basil ...
– typical bombardier beetles
Broscinae
Broscinae is a subfamily of 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 A ...
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
* '' Coarazup ...
(sometimes in Harpalinae)
Gineminae
''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 sp ...
(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 apomorphi ...
(over 20,000 species)
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 (sometimes in Harpalinae)
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
* ''Ana ...
– including Chlaeniinae, Oodinae (sometimes in Harpalinae)
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
** '' Cymbionotu ...
Migadopinae
The Migadopinae are 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 ...
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 2 ...
(sometimes in Harpalinae)
Panagaeinae
Panagaeinae is a subfamily of beetles in the family Carabidae, containing the following genera:
* '' Agonica'' Sloane, 1920
* '' Ardistomopsis'' Straneo & Ball, 1989
* '' Bascanus'' Peringuey, 1896
* ''Brachygnathus'' Perty, 1830
* '' Calatho ...
(sometimes in Harpalinae)
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, 1 ...
(sometimes in Harpalinae)
Pseudomorphinae
Pseudomorphinae is a subfamily of ground beetles (family Carabidae). Occasionally it was treated as a tribe Pseudomorphini of subfamily Harpalinae
Harpalinae is a huge subfamily of ground beetles that contains 20,000 species or ~6,400 spp. ...
(sometimes in Harpalinae)
Psydrinae
Psydrinae is a subfamily of beetles in the family Carabidae.
Genera
The subfamily contains the following genera:
* ''Amblytelus'' Erichson, 1842
* ''Bembidiomorphum'' Champion, 1918
* ''Celanida'' Castelnau, 1867
* '' Dystrichothorax'' Blackbur ...
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. Ho ...
– 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'' Belouso ...
– 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
Introduction
Amarotypines are a small tribe of ground beetles ( Carabidae), 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 a ...
– Migadopinae or a distinct subfamily?
* Gehringiini – Psydrinae, Trechinae, or a distinct subfamily?
Notes
References
Further reading
*
*
*
*
External links
Carabidae of the World
{{DEFAULTSORT:Ground Beetle