Ground beetle: Wikis

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Ground beetles
Golden Ground Beetle (Carabus auratus: Carabinae, Carabini) predating an earthworm
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Suborder: Adephaga
Superfamily: Caraboidea
Family: Carabidae
Latreille, 1802
Subfamilies

Amblytelinae (disputed)
Apotominae
Brachininae
Broscinae
Carabinae
Cicindelinae (disputed)
Cicindinae
Elaphrinae
Harpalinae
Hiletinae
Loricerinae
Melaeninae
Migadopinae
Nebriinae (disputed)
Nototylinae
Omophroninae
Paussinae
Promecognathinae
Psydrinae
Scaritinae
Siagoninae
Trechinae
and see text

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.[1]

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[2].

A Brachinus sp. typical bombardier beetle (Brachininae: Brachinini) from North Carolina
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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.[3]

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,[4] but this is based on a misreading of his autobiography[5]; 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!"[6]

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 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.'"[7]

Evolution and systematics

The Adephaga 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.[8]

Subfamilies and selected genera

Carabus violaceus (Carabinae: Carabini)

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 the Harpalinae instead of lumping them, restricting them to what in the system use 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. 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

Elaphrus cupreus (Elaphrinae: Elaphrini)
Loricera pilicornis (Loricerinae: Loricerini)
Notiophilus palustris (Nebriinae: Notiophilini)
Clivina fossor (Scaritinae: Clivinini)

Carabinae Latreille, 1802 - including Agoninae, Callistinae

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

Cicindinae

Elaphrinae Latreille, 1802

Hiletinae

Loricerinae Bonelli, 1810

Migadopinae

Nebriinae (includes Notiophilinae, often included in Carabinae)

Nototylinae

Omophroninae Bonelli, 1810 – round sand beetles

Paussinae Latreille, 1807 – ant nest beetles, flanged bombardier beetles (c.50 genera)

Promecognathinae

Scaritinae Bonelli, 1810 – pedunculate ground beetles

  • Clivina Latreille, 1802
  • Dyschirius Bonelli, 1810
  • Scarites Fabricius, 1775

Siagoninae Bonelli, 1810

Carabidae Conjunctae

Amblytelus sp. from Australia (Amblytelinae: Amblytelini)
Broscus cephalotes (Broscinae: Broscini)
Egyptian Predator Beetle, Anthia sexguttata (Harpalinae: Anthiini)
Mormolyce phyllodes (Harpalinae: Lebiini)
Bembidion quadrimaculatum (Trechinae: Bembidiini)

Amblytelinae[14] Sloane, 1898

Apotominae

  • Apotomus Illiger, 1807

Brachininae Bonelli, 1810

  • Aptinus Bonelli, 1810 (tentatively placed here)
  • Brachinus Weber, 1801
  • 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 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
  • Mecodema
  • 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)

Melaeninae

Psydrinae

  • Mecyclothorax

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)
  • Pseudanophthalmus Jeannel, 1920
  • 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

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


Footnotes

  1. ^ Kromp (1999)
  2. ^ Encyclopedia of Entomology by John L. Capinera pg.1746
  3. ^ Huey & Pianka (1977)
  4. ^ AMNH (2005)
  5. ^ Barlow (1958): p.62
  6. ^ Darwin (1846)
  7. ^ Barlow (1958): p.63-64
  8. ^ Molecular Phylogeny and Evolution of Carabid Ground Beetles By Syozo Osawa
  9. ^ Löbl & Smetana (2003-)
  10. ^ CWD (2008)
  11. ^ Maddison (1995)
  12. ^ Maddison (2006)
  13. ^ FEWS (2004)
  14. ^ 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: Maddison (1999)

References

External Links


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