Ground beetles are a large, cosmopolitan family of beetles, Carabidae, with more than 40,000
Ground beetles are a large, cosmopolitan family of beetles, Carabidae, with more than 40,000

species worldwide, approximately 2,000 of which are found in North America and 2,700
species worldwide, approximately 2,000 of which are found in North America and 2,700

in Europe.
in Europe.

Description and ecology Although their body shapes and coloring vary
Description and ecology Although their body shapes and coloring vary

somewhat, most are shiny black or metallic and have ridged wing covers. The elytra are
somewhat, most are shiny black or metallic and have ridged wing covers. The elytra are

fused in some species, particularly large Carabinae, rendering the beetles unable to
fused in some species, particularly large Carabinae, rendering the beetles unable to

fly. The genus Mormolyce is known as violin beetles due to their peculiarly shaped elytra.
fly. The genus Mormolyce is known as violin beetles due to their peculiarly shaped elytra.

All carabids except the quite primitive flanged bombardier beetles have a groove on their
All carabids except the quite primitive flanged bombardier beetles have a groove on their

foreleg tibiae bearing a comb of hairs used for cleaning their antennae.
foreleg tibiae bearing a comb of hairs used for cleaning their antennae.

Defensive secretions Typical for the ancient beetle suborder Adephaga
Defensive secretions Typical for the ancient beetle suborder Adephaga

to which they belong, they have paired pygidial glands in the lower back of the abdomen. These
to which they belong, they have paired pygidial glands in the lower back of the abdomen. These

are well developed in ground beetles, and produce noxious or even caustic secretions
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 beetles, these
used to deter would-be predators. In some, commonly known as bombardier beetles, these

secretions are mixed with volatile compounds and ejected by a small combustion, producing
secretions are mixed with volatile compounds and ejected by a small combustion, producing

a loud popping sound and a cloud of hot and acrid gas which can injure small mammals,
a loud popping sound and a cloud of hot and acrid gas which can injure small mammals,

such as shrews, and is liable to kill invertebrate predators outright. To humans, getting "bombed"
such as shrews, and is liable to kill invertebrate predators outright. To humans, getting "bombed"

by a bombardier beetle is a decidedly unpleasant experience. This ability has evolved independently
by a bombardier beetle is a decidedly unpleasant experience. This ability has evolved independently

twice as it seems – in the flanged bombardier beetles which are among the most ancient ground
twice as it seems – in the flanged bombardier beetles which are among the most ancient ground

beetles, as well as in the typical bombardier beetles which are part of a more "modern"
beetles, as well as in the typical bombardier beetles which are part of a more "modern"

lineage. The Anthiini, meanwhile, can mechanically squirt their defensive secretions for considerable
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 they are
distances and are able to aim with a startling degree of accuracy; in Afrikaans they are

known as oogpisters. In one of the very few known cases of a vertebrate mimicking an arthropod,
known as oogpisters. In one of the very few known cases of a vertebrate mimicking an arthropod,

juvenile Heliobolus lugubris lizards are colored similar to the aposematic oogpister beetles,
juvenile Heliobolus lugubris lizards are colored similar to the aposematic oogpister beetles,

and move in a way that makes them look surprisingly similar to the insects at a casual glance.
and move in a way that makes them look surprisingly similar to the insects at a casual glance.

Charles Darwin may have found himself on the receiving end of a bombardier beetle's defences
Charles Darwin may have 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;
on a collecting trip in 1828, but this is based on a misreading of his autobiography;

a bombardier beetle's "bombing" is already triggered by picking it up, and Darwin had
a bombardier beetle's "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
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
ran afoul of its secretions. He discussed this incident and another such case in a letter

to Leonard Jenyns as follows:
to Leonard Jenyns as follows:

A Cychrus rostratus once squirted into my eye & gave me extreme pain; & I must tell
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 in my early entomological days; under
you what happened to me on the banks of the Cam in my early entomological days; under

a piece of bark I found two carabi & caught one in each hand, when lo & behold I saw a
a piece of bark I found two carabi & 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
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
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
between my teeth, when to my unspeakable disgust & pain the little inconsiderate beast squirted

his acid down my throat & I lost both Carabi & Panagæus!
his acid down my throat & I lost both Carabi & Panagæus!

Ecology Common habitats are under the bark of trees,
Ecology Common habitats are under the bark of trees,

under logs, or among rocks or sand by the edge of ponds and rivers. Most species are
under logs, or among rocks or sand by the edge of ponds and rivers. Most species are

carnivorous and actively hunt for any invertebrate prey they can overpower. Some will run swiftly
carnivorous and actively hunt for any invertebrate prey they can overpower. Some will run swiftly

to catch their prey; tiger beetles can sustain speeds of 8 km/h – in relation to their
to catch their prey; tiger beetles can sustain speeds of 8 km/h – in relation to their

body length they are among the fastest land animals on Earth. Unlike most Carabidae which
body length they are among the fastest land animals on Earth. Unlike most Carabidae which

are nocturnal, the tiger beetles are active diurnal hunters and often brightly coloured;
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
they have large eyes and hunt by sight. Ground beetles of the species Promecognathus laevissimus

are specialised predators of the cyanide millipede Harpaphe haydeniana, countering the hydrogen
are specialised predators of the cyanide millipede Harpaphe haydeniana, countering the hydrogen

cyanide which makes these millipedes poisonous to most carnivores.
cyanide which makes these millipedes poisonous to most carnivores.

Relationship with humans As predators of invertebrates, including many
Relationship with humans As predators of invertebrates, including many

pests, most ground beetles are considered beneficial organisms. The caterpillar hunters
pests, most ground beetles are considered beneficial organisms. The caterpillar hunters

are famous for their habit of devouring prey in quantity, eagerly feeding on tussock moth
are famous for their habit of devouring prey in quantity, eagerly feeding on tussock moth

caterpillars, processionary caterpillars and woolly worms, which due to their urticating
caterpillars, processionary caterpillars and woolly worms, which due to their urticating

hairs are avoided by most insectivores. Large numbers of the Forest Caterpillar Hunter,
hairs are avoided by most insectivores. Large numbers of the Forest Caterpillar Hunter,

native to Europe, were shipped to New England for biological control of the gypsy moth as
native to Europe, were shipped to New England for biological control of the gypsy moth as

early as 1905. A few species are nuisance pests. Zabrus is
early as 1905. A few species are nuisance pests. Zabrus is

one of the few herbivorous ground beetle genera, and on rare occasions Zabrus tenebrioides
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 crops. Large species,
for example occurs abundantly enough to cause some damage to grain crops. Large species,

usually Carabinae, can become a nuisance if present in numbers, particularly during outdoor
usually Carabinae, can become a nuisance if present in numbers, particularly during outdoor

activities such as camping; they will void their defensive secretions when threatened,
activities such as camping; they will void their defensive secretions when threatened,

and if they hide among provisions this can despoil food. Since ground beetles are generally
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
reluctant or even unable to fly, it is usually easy to block their potential routes of entry

mechanically or with a topical insecticide.
mechanically or with a topical insecticide.

Especially in the 19th century and to a lesser extent today, their large size and conspicuous
Especially in the 19th century and to a lesser extent today, their large size and conspicuous

coloration as well as the odd morphology of some made many ground beetles a popular object
coloration as well as the odd morphology of some made many ground beetles a popular object

of collection and study for professional and amateur coleopterologists. High prices were
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
paid for rare and exotic specimens, and in the early to mid-19th century there was a

veritable "beetle craze" in England. As mentioned above, Charles Darwin was an ardent collector
veritable "beetle craze" in England. As mentioned above, Charles Darwin was an ardent collector

of beetles when he was about twenty years old, to the extent that he'd rather scour
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, John Stevens Henslow and Henry
the countryside for rare specimens with William Darwin Fox, John Stevens Henslow and Henry

Thompson than to study theology as his father wanted him to do. In his autobiography he
Thompson than to study theology as his father wanted him to do. In his autobiography he

fondly recalled his experiences with Licinus and Panagaeus, and wrote:
fondly recalled his experiences with Licinus and Panagaeus, and wrote:

No poet ever felt more delight at seeing his first poem published than I did at seeing
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."
in Stephen's Illustrations of British Insects the magic words, "captured by C. Darwin, Esq."

Evolution and systematics The Adephaga are documented since the end
Evolution and systematics The Adephaga are documented since the end

of the Permian, about 250 million years ago. Ground beetles evolved in the latter Triassic,
of the Permian, about 250 million years ago. Ground beetles evolved in the latter Triassic,

having separated from their closest relatives by 200 million years ago. The family diversified
having separated from their closest relatives by 200 million years ago. The family diversified

throughout the Jurassic, and the more advanced lineages, such as the Harpalinae, underwent
throughout the Jurassic, and the more advanced lineages, such as the Harpalinae, underwent

a vigorous radiation starting in the Cretaceous. The closest living relatives of the ground
a vigorous radiation starting in the Cretaceous. The closest living relatives of the ground

beetles are the false ground beetles and the wrinkled bark beetles. They are sometimes
beetles are the false ground beetles and the wrinkled bark beetles. They are sometimes

even included in the Carabidae as subfamilies or as tribes incertae sedis, but more preferably
even included in the Carabidae as subfamilies or as tribes incertae sedis, but more preferably

they are united with the ground beetles in the superfamily Caraboidea.
they are united with the ground beetles in the superfamily Caraboidea.

Much research has been done on elucidating the phylogeny of the ground beetles and adjusting
Much research has been done on elucidating the phylogeny of the ground beetles and adjusting

systematics and taxonomy accordingly. While there is no completely firm consensus, a few
systematics and taxonomy 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
points are generally accepted: As it seems, the ground beetles consist of a number of

more basal lineages and the extremely diverse Harpalinae which contain over half the described
more basal lineages and the extremely diverse Harpalinae which contain over half the described

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

Subfamilies and selected genera The taxonomy used here is based on the Catalogue
Subfamilies and selected genera The taxonomy used here is based on the Catalogue

of Palaearctic Coleoptera and the Carabidae of the World Database. Other classifications,
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
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.
and the more advanced group informally called "Carabidae Conjunctae", differ in details.

For example, the system used by the Tree of Life Web Project makes little use of subfamilies,
For example, the system used by the Tree of Life Web Project makes little use of subfamilies,

listing most tribes as incertae sedis as to subfamily. Fauna Europaea on the other hand
listing most tribes as incertae sedis as to subfamily. Fauna Europaea on the other hand

splits rather than lumps the Harpalinae, restricting them to what in the system used here is the
splits rather than lumps the Harpalinae, restricting them to what in the system used here is the

tribe Harpalini. All the approaches mentioned above are legitimate
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
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
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.
majority of authors presently uses this system, following the Catalogue of Palaearctic Coleoptera.

Second, the MediaWiki markup cannot at present adequately represent the relationships of
Second, the MediaWiki markup cannot at present adequately represent the relationships of

the ground beetle subgroups in detail if the restricted view of the Harpalinae is chosen.
the ground beetle subgroups in detail if the restricted view of the Harpalinae is chosen.

Basal ground beetles
Basal ground beetles

Carabinae Latreille, 1802 – including Agoninae and Callistinae
Carabinae Latreille, 1802 – including Agoninae and Callistinae

Altagonum Aplothorax
Altagonum Aplothorax

Calosoma - including Callisthenes Carabus
Calosoma - including Callisthenes Carabus

Cychrus Homothes
Cychrus Homothes

Fortagonum Laemostenus
Fortagonum Laemostenus

Notagonum Cicindelinae – tiger beetles
Notagonum Cicindelinae – tiger beetles

Cicindinae Elaphrinae Latreille, 1802
Cicindinae Elaphrinae Latreille, 1802

Blethisa Diacheila Motschulsky, 1844
Blethisa Diacheila Motschulsky, 1844

Elaphrus Hiletinae
Elaphrus Hiletinae

Loricerinae Bonelli, 1810 Loricera
Loricerinae Bonelli, 1810 Loricera

Migadopinae Nebriinae
Migadopinae Nebriinae

Leistus Nebria
Leistus Nebria

Notiophilus Pelophila Dejean, 1821
Notiophilus Pelophila Dejean, 1821

Nototylinae Omophroninae Bonelli, 1810 – round sand
Nototylinae Omophroninae Bonelli, 1810 – round sand

beetles Omophron
beetles Omophron

Paussinae – ant nest beetles, flanged bombardier beetles
Paussinae – ant nest beetles, flanged bombardier beetles

Promecognathinae Scaritinae Bonelli, 1810 – pedunculate ground
Promecognathinae Scaritinae Bonelli, 1810 – pedunculate ground

beetles Clivina
beetles Clivina

Dyschirius Bonelli, 1810 Scarites
Dyschirius Bonelli, 1810 Scarites

Siagoninae Bonelli, 1810 Carabidae Conjunctae
Siagoninae Bonelli, 1810 Carabidae Conjunctae

Amblytelinae Sloane, 1898 Amblytelus
Amblytelinae Sloane, 1898 Amblytelus

Apotominae Brachininae – typical bombardier beetles
Apotominae Brachininae – typical bombardier beetles

Broscinae Dryptinae
Broscinae Dryptinae

Gineminae Harpalinae
Gineminae Harpalinae

Lebiinae – including Cyclosominae, Mormolycinae, Odacanthinae, Perigoninae
Lebiinae – including Cyclosominae, Mormolycinae, Odacanthinae, Perigoninae

Licininae – including Chlaeniinae, Oodinae Melaeninae
Licininae – including Chlaeniinae, Oodinae Melaeninae

Migadopinae Orthogoniinae
Migadopinae Orthogoniinae

Panagaeinae Platyninae
Panagaeinae Platyninae

Pseudomorphinae Psydrinae
Pseudomorphinae Psydrinae

Pterostichinae – including Zabrinae Trechinae – including Bembidiinae, Patrobinae
Pterostichinae – including Zabrinae Trechinae – including Bembidiinae, Patrobinae

Tribes incertae sedis Amarotypini – Migadopinae or a distinct
Tribes incertae sedis Amarotypini – Migadopinae or a distinct

subfamily? Gehringiini – Psydrinae, Trechinae or a
subfamily? Gehringiini – Psydrinae, Trechinae or a

distinct subfamily? Notes
distinct subfamily? Notes

References
References

Further reading
Further reading

E. Csiki. Die Käferfauna des Karpaten-Beckens [The beetle fauna of the Carparthian basin].
E. Csiki. Die Käferfauna des Karpaten-Beckens [The beetle fauna of the Carparthian basin].

Budapest. pp. 71–546.  K. Kult. Klíč k určování brouků čeledi
Budapest. pp. 71–546.  K. Kult. Klíč k určování brouků čeledi

Carabidae Československé republiky [Key to the beetles of family Carabidae of the
Carabidae Československé republiky [Key to the beetles of family Carabidae of the

Czech Republic]. Prague.  C. H. Lindroth. Coleoptera, Carabidae. Svensk
Czech Republic]. Prague.  C. H. Lindroth. Coleoptera, Carabidae. Svensk

Insectenfauna, Vol. 9. Stockholm. pp. 1–260.  Edmund Reitter. Die Käfer des Deutschen Reiches
Insectenfauna, Vol. 9. Stockholm. pp. 1–260.  Edmund Reitter. Die Käfer des Deutschen Reiches

[The beetles of the German Empire]. Stuttgart: K. G. Lutz. 
[The beetles of the German Empire]. Stuttgart: K. G. Lutz. 

External links Carabidae of the World
External links Carabidae of the World