Ground beetle

Ground beetle
Ground beetles
Golden ground beetle eating an earthworm
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Suborder: Adephaga
(unranked): Geadephaga
Superfamily: Caraboidea
Family: Carabidae
Latreille, 1802
Subfamilies[1]

Anthiinae
Apotominae
Brachininae
Broscinae
Carabinae
Cicindelinae
Ctenodactylinae
Dryptinae
Elaphrinae
Gineminae
Harpalinae
Hiletinae
Lebiinae
Licininae
Loricerinae
Melaeninae
Migadopinae
Nebriinae
Nototylinae
Omophroninae
Orthogoniinae
Panagaeinae
Paussinae
Platyninae
Promecognathinae
Protorabinae
Pseudomorphinae
Psydrinae
Pterostichinae
Rhysodinae
Scaritinae
Siagoninae
Trechinae
Xenaroswellianinae
(See text)

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 in Europe.[2]

Contents

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 (Paussinae) have a groove on their foreleg tibiae bearing a comb of hairs. This is used for cleaning their antennae.[3]

A Brachinus sp. typical bombardier beetle (Brachininae: Brachinini) from North Carolina

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 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 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 like shrews, 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 (Paussinae) which are among the most ancient ground beetles, as well as in the typical bombardier beetles (Brachininae) 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 an arthropod, 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.[4]

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,[5] but this is based on a misreading of his autobiography;[6][not in citation given] 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 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!"[7]

A Lophyra sp. tiger beetle from Tanzania

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 carnivorous and 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 Harpaphe haydeniana, 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 (Calosoma) are famous for their habit of devouring insect larvae and pupae 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 (C. sycophanta), native to Europe, were shipped to New England for biological control of the Gypsy Moth (Lymantria dispar) 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 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 insecticide.

A crucifix ground beetle (Panagaeus cruxmajor) got Charles Darwin into trouble in 1828.

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 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 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.'"[6]

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, 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 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.[8]

Subfamilies and selected genera

The taxonomy used here is based on the Catalogue of Palaearctic Coleoptera[9] and the Carabidae of the World Database.[10] 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",[11] 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.[12] Fauna Europaea on the other hand splits rather than lumps the Harpalinae, restricting them to what in the system uses here is the tribe Harpalini.[13]

All the approaches mentioned above are legitimate as they agree with the phylogeny as far as it has been resolved.[citation needed] 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.[citation needed]

Basal ground beetles

Carabus lateralis (Carabinae: Carabini)
Loricera pilicornis (Loricerinae: Loricerini)
Notiophilus substriatus (Nebriinae: Notiophilini)
Clivina fossor (Scaritinae: Clivinini)
Amblytelus (Amblytelinae: Amblytelini)

Carabinae Latreille, 1802 – including Agoninae and Callistinae

  • Altagonum
  • Aplothorax (monotypic genus)
  • Calosoma - including Callisthenes
  • Carabus
  • Cychrus
  • Homothes
  • Fortagonum
  • Laemostenus
  • Notagonum

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

Cicindinae

Elaphrinae Latreille, 1802

Hiletinae

Loricerinae Bonelli, 1810

  • Loricera

Migadopinae

Nebriinae (includes Notiophilinae, often included in Carabinae)

Nototylinae

Omophroninae Bonelli, 1810 – round sand beetles

Paussinae – ant nest beetles, flanged bombardier beetles

Promecognathinae

Scaritinae Bonelli, 1810 – pedunculate ground beetles

  • Clivina
  • Dyschirius Bonelli, 1810
  • Scarites

Siagoninae Bonelli, 1810

Carabidae Conjunctae

Broscus cephalotes (Broscinae: Broscini)
Dixus sphaerocephalus (Harpalinae: Harpalini)
Mormolyce phyllodes (Harpalinae: Lebiini)
Bembidion quadrimaculatum (Trechinae: Bembidiini)

Amblytelinae Sloane, 1898[Note 1]

  • Amblytelus

Apotominae

  • Apotomus Illiger, 1807

Brachininae Bonelli, 1810 – typical bombardier beetles

  • Aptinus (tentatively placed here)
  • Brachinus
  • Mastax Fischer von Waldheim, 1828

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
  • Brullea Laporte de Castelnau, 1868
  • Cascellius Curtis, 1839
  • 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)

Melaeninae

Psydrinae

  • Mecyclothorax

Trechinae Bonelli, 1810 – including Bembidiinae, Patrobinae

  • Aepus Samouelle, 1819
  • Amerizus de Chaudoir 1868 – including Gnatholymnaeum
  • Anillinus Casey, 1918
  • Anophthalmus
  • Asaphidion
  • Bembidion
  • Blemus – including Lasiotrechus
  • Cardiaderus Dejean, 1828
  • Cillenus Leach, 1819
  • Deltomerus Motschulsky, 1850
  • Duvaliopsis Jeannel, 1928 (tentatively placed here)
  • Duvalius Delarouzée, 1859
  • Lymnastis Motschulsky, 1862
  • Miscodera Eschscholtz, 1830
  • Ocys Stephens, 1828
  • Patrobus
  • Perileptus Schaum, 1860
  • Pogonus Dejean, 1821
  • Porotachys Netolitzky, 1914 (tentatively placed here)
  • Pseudanophthalmus
  • Pseudaphaenops Winkler, 1912
  • Serranillus Barr, 1996
  • Tachyra Motschulsky, 1862
  • Tachys Dejean, 1821
  • Tachyta Kirby, 1837
  • Thalassophilus Wollaston, 1854
  • Trechoblemus Ganglbauer, 1891
  • Trechosia Jeannel, 1926
  • Trechus
  • Winklerites Jeannel, 1937

Tribes incertae sedis

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

Notes

  1. ^ Usually placed in the Psydrinae or Trechinae, 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.[14]

References

  1. ^ "Carabidae Taxa". Carabidae of the World. 2011. http://carabidae.pro/carabidae. Retrieved 24 Jun 2011. 
  2. ^ B. Kromp (1999). "Carabid beetles in sustainable agriculture: a review on pest control efficacy, cultivation aspects and enhancement". Agriculture, Ecosystems and Environment 74 (1–3): 187–228. doi:10.1016/S0167-8809(99)00037-7. 
  3. ^ John L. Capinera. Encyclopedia of Entomology. p. 1746. 
  4. ^ R. B. Huey & E. R. Pianka (1977). "Natural selection for juvenile lizards mimicking noxious beetles". Science 195 (4274): 201–203. doi:10.1126/science.831272. PMID 831272. 
  5. ^ "Young Naturalist, A Lifelong Passion". Darwin. American Museum of Natural History. 2005. http://www.amnh.org/exhibitions/darwin/young/lifelong.php. Retrieved February 16, 2011. 
  6. ^ a b Nora Barlow, ed (1958). "Cambridge, 1828–1831". The Autobiography of Charles Darwin. pp. 56–71. http://darwin-online.org.uk/content/frameset?viewtype=side&itemID=F1497&pageseq=64. 
  7. ^ Charles Darwin (1846). "Letter to Leonard Jenyns, October 17, 1846". http://www.darwinproject.ac.uk/darwinletters/calendar/entry-1009.html. 
  8. ^ Shōzō Ōsawa, Zhi-Hui Su & Yūki Inmura (2004). Molecular Phylogeny and Evolution of Carabid Ground Beetles. Springer. ISBN 4-431-00487-4. 
  9. ^ I. Löbl & A. Smetana, ed (2003–). Catalogue of Palaearctic Coleoptera. Stenstrup, Denmark: Apollo Books. 
  10. ^ "Trees of family Carabidae". Carabidae of the World Database. 2008. http://www.carabidae.ru/carabidae.htm. Retrieved July 24, 2008. 
  11. ^ David R. Maddison (January 1, 1995). "Carabidae Conjunctae". Tree of Life Web Project. http://tolweb.org/Carabidae_Conjunctae/66/1995.01.01. Retrieved July 24, 2008. 
  12. ^ David R. Maddison (April 11, 2006). "Carabidae. Ground beetles and tiger beetles". Tree of Life Web Project. http://tolweb.org/Carabidae/8895/2006.04.11. Retrieved July 24, 2008. 
  13. ^ "Harpalinae". Fauna Europaea. 2004. http://www.faunaeur.org/full_results.php?id=379457. Retrieved February 16, 2011. 
  14. ^ David R. Maddison (January 1, 1999). "Amblytelini". Tree of Life Web Project. http://tolweb.org/Amblytelini/97/1999.01.01. Retrieved July 24, 2008. 

Further reading

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  • E. Csiki (1946) (in German). Die Käferfauna des Karpaten-Beckens [The beetle fauna of the Carparthian basin]. Budapest. pp. 71–546. 
  • K. Kult (1947) (in Czech). Klíč k určování brouků čeledi Carabidae Československé republiky [Key to the beetles of family Carabidae of the Czech Republic]. Prague. 
  • C. H. Lindroth (1942) (in Swedish). Coleoptera, Carabidae. Svensk Insectenfauna, Vol. 9. Stockholm. pp. 1–260. 
  • Edmund Reitter (1908–1917) (in German). Die Käfer des Deutschen Reiches [The beetles of the German Empire]. Stuttgart: K. G. Lutz. 

External links


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Look at other dictionaries:

  • Ground beetle — ground ground (ground), n. [OE. ground, grund, AS. grund; akin to D. grond, OS., G., Sw., & Dan. grund, Icel. grunnr bottom, Goth. grundus (in composition); perh. orig. meaning, dust, gravel, and if so perh. akin to E. grind.] 1. The surface of… …   The Collaborative International Dictionary of English

  • ground-beetle — groundˈ beetle noun Any beetle of the Carabidae, a family related to the tiger beetles • • • Main Entry: ↑ground …   Useful english dictionary

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  • ground beetle — any of numerous nocturnal, terrestrial beetles of the family Carabidae that feed chiefly on other insects. [1840 50] * * * ▪ insect       any member of over 30,000 insect species in one of the largest families in the insect order Coleoptera. They …   Universalium

  • ground beetle — ground′ bee tle n. ent any of numerous nocturnal, terrestrial beetles of the family Carabidae that feed chiefly on other insects • Etymology: 1840–50 …   From formal English to slang

  • ground beetle — noun predacious shining black or metallic terrestrial beetle that destroys many injurious insects • Syn: ↑carabid beetle • Hypernyms: ↑beetle • Hyponyms: ↑bombardier beetle, ↑calosoma • Member Holonyms: ↑ …   Useful english dictionary

  • ground beetle — noun Date: 1848 any of a large cosmopolitan family (Carabidae) of soil inhabiting usually carnivorous often shiny black or metallic beetles commonly having fused elytra …   New Collegiate Dictionary

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  • List of ground beetle (Carabidae) species recorded in Britain — The following is a list of the ground beetles recorded in Britain, organised by subfamily ( inae enddings) and by tribe ( ini endings). For other beetle families, see the main list of beetle species recorded in Britain.Cicindelinae* Cicindela… …   Wikipedia

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