Ground beetles or carabids are collective terms for the beetle family Carabidae. This is a large family, with more than 40,000 species worldwide, approximately 2,000 of which are found in North America and 2,700 in Europe.
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
, particularly large Carabinae
, rendering the beetles unable to 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 foreleg tibiae
bearing a comb of hairs. This is used for cleaning their antennae
. However, rather than the flanged bombardier beetles never evolving this comb, it seems rather that they lost it, demonstrating that evolution
does not equal progress
but merely change.
Typical for the ancient beetle suborder Adephaga
to which they belong, they have paired pygidial glands
on the lower back of the abdomen
. These produce noxious or even caustic
secretions 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 a loud popping sound and a cloud of hot and acrid gas which can injure small mammals
, and is liable to kill invertebrate
predators outright. To humans, getting "bombed" by a bombardier beetle is a decidedly unpleasant experience. The "bombing" ability has evolved
independently 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" 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 known as oogpisters
("eye-pissers"). In one of the very few known cases of a vertebrate mimicking
, 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.
It is sometimes suggested that Charles Darwin 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; 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 when 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 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!
are under the bark of trees, under logs, or among rocks or sand by the edge of ponds and rivers. Most species are carnivorous
actively hunt for any invertebrate
prey they can overpower. Some will run swiftly to catch their prey; tiger beetles
(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
), countering the hydrogen cyanide
which makes these millipedes poisonous to most carnivores.
Relationship with humans
As predators of invertebrates
, including many pests
, most ground beetles are considered beneficial organisms. The caterpillar hunters
) are famous for their habit of devouring insect larvae
in quantity, eagerly feeding on tussock moth
(Lymantriidae) caterpillars, processionary caterpillars
(Thaumetopoeidae) and woolly worms
(Arctiidae), which due to their urticating 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 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 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 activities such as camping; they will void their defensive secretions when threatened and if they hide between 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 insecticide.
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 (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. As mentioned above, Charles Darwin was an ardent collector of beetles around when he was about twenty years old, to the extent that he rather went out to hunt for rare specimens with William Darwin Fox, John Stevens Henslow and Sir Henry 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:
"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
are documented since the end of the Permian
, about 250 million years ago
(mya). Ground beetles evolved
in the latter Triassic
, having separated from their closest relatives by 200 mya. The family diversified 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 beetles are the false ground beetles
(Trachypachidae) and the wrinkled bark beetles
(Rhysodidae). They are sometimes 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
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 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 species and into which several formerly independent families had to be subsumed.
Subfamilies and selected genera
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
makes little use of subfamilies, listing most tribes
as incertae sedis
as to subfamily. Fauna Europaea
on the other hand splits the Harpalinae instead of lumping them
, restricting them to what in the system use 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 markup cannot at present adequately represent the relationships of the ground beetle subgroups in detail if the restricted view of the Harpalinae is chosen.
Basal ground beetles
Carabinae Latreille, 1802
- including Agoninae, Callistinae
Cicindelinae – tiger beetles (roughly 2,100 species); sometimes included in Carabidae
Elaphrinae Latreille, 1802
Loricerinae Bonelli, 1810
Nebriinae - includes Notiophilinae, often included in Carabinae
Omophroninae Bonelli, 1810 – round sand beetles
Paussinae Latreille, 1807 – flanged bombardier beetles
Scaritinae Bonelli, 1810 – pedunculate ground beetles
Siagoninae Bonelli, 1810
Amblytelinae Sloane, 1898 - usually placed in the Psydrinae or Trechinae, but they seem to represent a distinct lineage related to Brachininae and Harpalinae, and in the system used here would consequently be eligible for subfamily status.
Brachininae Bonelli, 1810
Broscinae Hope, 1838
- Acallistus Sharp, 1886
- Adotela Laporte de Castelnau, 1867
- Axonya Andrewes, 1923
- Baripus Dejean, 1828
- Bountya Townsend, 1971
- Brithysternum Macleay, 1873
- Broscodera Lindroth, 1961
- Broscodes Bolivar, 1914
- Broscosoma Rosenhauer, 1846
- Broscus Panzer, 1813
- Brullea Laporte de Castelnau, 1868
- Cascellius Curtis, 1839
- Cerotalis Laporte de Castelnau, 1868
- Chaetobroscus Semenov, 1900
- Chylnus Sloane, 1920
- Craspedonotus Schaum, 1863
- Creobius Guérin-Méneville, 1838
- Diglymma Sharp, 1886
- Eobroscus Kruizhanovskií, 1951
- Eurylychnus Bates, 1891
- Gnathoxys Westwood, 1842
- Metaglymma Bates, 1867
- Microbarypus Roig-Juñent, 2000
- Miscodera Eschscholtz, 1830
- Nothobroscus Roig-Juñent & Ball, 1995
- Nothocascellius Roig-Juñent, 1995
- Oregus Putzeys, 1868
- Parroa Laporte de Castelnau, 1868
- Percolestus Sloane, 1892
- Percosoma Schaum, 1858
- Promecoderus Dejean, 1829
- Rawlinsius Davidson & Ball, 1998
- Zacotus Leconte, 1869
Harpalinae - including Chlaeniinae, Cyclosominae, Dryptinae, Lebiinae, Licininae, Mormolycinae, Odacanthinae, Oodinae, Panagaeinae, Perigoninae, Platyninae, Pseudomorphinae, Pterostichinae, Zabrinae (over 20,000 species)
Trechinae Bonelli, 1810 - including Bembidiinae, Patrobinae
- Aepus Samouelle, 1819
- Amerizus de Chaudoir 1868 - includes Gnatholymnaeum
- Anillinus Casey, 1918
- Anophthalmus Sturm, 1844
- Asaphidion Des Gozis, 1886
- Bembidion Latreille, 1802
- Blemus Dejean, 1821 - including Lasiotrechus
- Broscus Panzer, 1813
- Cardiaderus Dejean, 1828
- Cillenus Leach, 1819
- Deltomerus Motschulsky, 1850
- Duvaliopsis Jeannel, 1928 (tentatively placed here)
- Duvalius Delarouzée, 1859
- Eurytrachelus Motschulsky, 1850
- Lymnastis Motschulsky, 1862
- Miscodera Eschscholtz, 1830
- Ocys Stephens, 1828
- Patrobus Dejean, 1821
- Perileptus Schaum, 1860
- Pogonus Dejean, 1821
- Porotachys Netolitzky, 1914 (tentatively placed here)
- Pseudaphaenops Winkler, 1912
- Rhysodes Dejean, 1821
- Serranillus Barr, 1996
- Tachyra Motschulsky, 1862
- Tachys Dejean, 1821
- Tachyta Kirby, 1837
- Thalassophilus Wollaston, 1854
- Trechoblemus Ganglbauer, 1891
- Trechosia Jeannel, 1926
- Trechus Clairville, 1806
Tribes incertae sedis
- (2005): Darwin - Young Naturalist, A Lifelong Passion Retrieved 2008-JUL-24.
- (1958): The autobiography of Charles Darwin 1809-1882. With the original omissions restored. Edited and with appendix and notes by his granddaughter Nora Barlow. Collins, London. HTML fulltext
- (2008): Trees of family Carabidae Retrieved 2008-JUL-24.
- (1946): Die Käferfauna des Karpaten-Beckens ["The beetle fauna of the Carparthian basin"] (vol. 1): 71-546. Budapest.
- (1846): [Letter to Leonard Jenyns, October 17 1846]. HTML fulltext
- (2004): Fauna Europaea version 1.1
- (1977): Natural selection for juvenile lizards mimicking noxious beetles. Science 195(4274): 201-203. (HTML abstract)
- (1999): Carabid beetles in sustainable agriculture: a review on pest control efficacy, cultivation aspects and enhancement. Agriculture, Ecosystems and Environment 74: 187-228.
- (1947): Klíč k určování brouků čeledi Carabidae Československé republiky ["Key to the beetles of family Carabidae of the Czech Republic"]. Prague.
- (1942): Coleoptera, Carabidae. In: Svensk Insectenfauna (Vol. 9): 1-260. Stockholm.
- (2003-): Catalogue of Palaearctic Coleoptera. Apollo Books, Stenstrup, Denmark.
- (1995): Tree of Life Web Project - Carabidae Conjunctae Version of 1995-JAN-01. Retrieved 2008-JUL-24.
- (1999): Tree of Life Web Project - Amblytelini Version of 1999-JAN-01. Retrieved 2008-JUL-24.
- (2006): Tree of Life Web Project - Carabidae. Ground beetles and tiger beetles Version of 2006-APR-11. Retrieved 2008-JUL-24.
- (1908-1917): Die Käfer des Deutschen Reiches ["The beetles of the German Reich"]. K.G. Lutz, Stuttgart.