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Carabidae

(Family)

Overview

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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]

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 unpl easant 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 Panag?us crux major; I could not bear to give up either of my Carabi, & to lose Panag?us 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 & Panag?us!"[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 spee ds 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 colored; 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 avo ided 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 wi th 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 th e 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 classificat ions, 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 o ne 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

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 ? ant nest beetles, flanged bombardier beetles

Promecognathinae

Scaritinae Bonelli, 1810 ? pedunculate ground beetles

Siagoninae Bonelli, 1810

Carabidae Conjunctae

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

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

Tribes incertae sedis

  • Amarotypini ? Harpalinae?< /li>
  • Gehringiini ? Trechinae or a distinct subfamily
  • Metiini ? Harpalinae?
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 t he 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 w ith 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 carr ying 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 Panag?us crux major; I could not bear to give up either of my Carabi, & to lose Panag?us 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 & Panag?us!"[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 colored; they have large eyes and hunt by sight. Ground beetles of the species Promecognathus laevissimus are specialised predators of the cyanide millipede Harpaphe hayd eniana, 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 abund antly 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 Le biini) 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

[ Back to top ]

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 con sist 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]

Basa l 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

  • Blethisa
  • Diacheila Motschulsky, 1844
  • Elaphrus

Hiletinae

Loricerinae Bonelli, 1810

  • Loricera

Migadopinae

Nebriinae (includes Notiophilinae, often included in Carabinae)

  • Leistus
  • Nebria
  • Notiophilus
  • Pelophila Dejean, 1821

Nototylinae

Omophroninae Bonelli, 1810 ? round sand beetles

  • Omophron

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)
  • Br achinus
  • Mastax Fischer von Waldheim, 1828

Broscinae Hope, 1838

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

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]
es (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 colored; 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 t heir 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 u sually 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 ext ent 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 Pal aearctic 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 legitim ate 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 (Carabin ae: 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

  • Blethisa
  • Diacheila Motschulsky, 1844
  • E laphrus

Hiletinae

Loricerinae Bonelli, 1810

  • Loricera

Migadopinae

Nebriinae (includes Notiophilinae, often included in Carabinae)

  • Leistus
  • Nebria
  • Notiophilus
  • Pelophila Dejean, 1821

Nototylinae

Omophroninae Bonelli, 1810 ? round sand beetles

  • Omophron

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

Harpalinae ? including Chlaeniinae, Cyclosominae, Dryptinae, Lebiinae, Licininae, Mormolycinae, Odacanthinae, Oodinae, Panagaeinae, Perigoninae, Platyninae, Pseudomorphinae, Pterostichinae, Zabrinae (over 20,000 species)

Me laeninae

Psydrinae

  • Mecyclothorax

Trechinae Bonelli, 1810 ? including Bembidiinae, Patrobinae

Tribes incertae sedis

  • Amarotypini ? Harpalinae?
  • Gehringiini ? Trechinae or a distinct subfamily
  • Metiini ? Harpalinae?
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 g round 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 Panag?us crux major; I could not bear to give up either of my Carabi, & to lose Panag?us 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 incon siderate beast squirted his acid down my throat & I lost both Carabi & Panag?us!"[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 colored; 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 o f 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 (P anagaeus 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 res earch 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 La treille, 1802 ? including Agoninae and 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

Omophronin ae Bonelli, 1810 ? round sand beetles

Paussinae ? ant nest beetles, flanged bombardier beetles

Promecognathinae

Scaritinae Bonelli, 1810 ? pedunculate ground beetles

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]

Apotominae

Brachininae Bonelli, 1810 ? typical bombardier beetles

Broscinae Hope, 1838

Harpalinae ? including Chlaeniinae, Cyclosominae, Dryptinae, Lebiinae, Licininae, Mormolycinae, Odacanthinae, Oodinae, Panagaeinae, Perigoninae, Platyninae, Pseudomorphinae, Pterostichinae, Zabrinae (over 20,000 species)

Melaeninae

Psydrinae

Trechinae Bonelli, 1810 ? including Bembidiinae, Patrobinae

Tribes incertae sedis

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 Passio n". 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). "Lett er to Leonard Jenyns, October 17, 1846". http://www.darwinproject.ac.uk/darwinletters/calendar/entry-1009.html
  8. ^ Shozo Osawa, Zhi-Hui Su & Yuki Inmura (2004). Molecular Phylogeny and Evolution of Carabid Ground Beetles. Springer. ISBN 4-431-00487-4. 
  9. ^ I. L?bl & 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_res ults.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. 

Taxonomy

The Family Carabidae is further organized into finer groupings including:

Genera

Abacetus

[more]

Abacidus

[more]

Abax

Sand table is a term for using constrained sand for modeling or educational purposes. The original version of a sand table may be the abax used by early Greek students. [more]

Abroscelis

Abroscelis is a genus of beetles in the family Carabidae, containing the following species: [more]

Acanthoscelis

[more]

Acinopus

[more]

Acupalpus

Acupalpus is a genus of insect-eating beetle species. Its representatives are found across Europe and Asia. [more]

Adelotopus

[more]

Adriaphaenops

[more]

Aephnidius

[more]

Aepopsis

[more]

Aepus

Aepus is a genus of ground beetles of the subfamily istributed in France, on Canary Islands and the archipelago Madeira in Spain, Albania, UK, Ireland, Norway and Sweden. [more]

Agatus

[more]

Agelaea

Agelaea is a genus of beetles in the family Carabidae, containing the following species: [more]

Agonocheila

[more]

Agonoderus

[more]

Agonotrechus

[more]

Agonum

Agonum is a large genus of ground beetles in the subfamily Harpalinae, tribe . They are mid-sized to smallish beetles, typically with dark metallic hues ? often reddish or bronze, but sometimes black, green etc. [more]

Agostenus

[more]

Agra

Agra (English pronunciation: ; Hindi: ????, Urdu: ? ???), the erstwhile capital of Hindustan, is a city on the banks of the river Yamuna in the northern state of Uttar Pradesh, India, 363 kilometres (226 mi) west of state capital, Lucknow and 200 kilometres (124 mi) south from national capital New Delhi. With a population of 1,686,976 (2010 est.), it is one of the most populous cities in Uttar Pradesh and the 19th most populous in India. Because 80 percent of the city's sewage flows into Yamuna River, it is 20th most polluted city in India. Agra can also refer to the administrative district that has its headquarters in Agra city. [more]

Ainocolpodes

[more]

Albanotrechus

[more]

Allegrettia

[more]

Allotriopus

[more]

Altagonum

[more]

Amara

Amara, the sun beetles, are a large genus of carabid beetles, mostly holarctic, but a few species are neotropical or occurring in eastern Asia. [more]

Amaroschema

[more]

Amblycheila

Amblycheila is a genus of flightless, nocturnal tiger beetles. There are at least five species occurring sparsely over much of the central and western United States, and with a few more species found southwards into Mexico. [more]

Amblygnathus

[more]

Amblystomus

Amblystomus is a genus of beetles in the family Carabidae, containing the following species: [more]

Amblytelus

[more]

Ambylgnathus

[more]

Amerinus

Amerinus linearis is a species of beetle in the family Carabidae, the only species in the genus Amerinus. [more]

Amerizus

[more]

Ameroduvalius

[more]

Amphasia

Amphasia is a genus of beetles in the family Carabidae, containing the following species: [more]

Amphimenes

[more]

Anasis

Anasis howitti is a species of beetle in the family Carabidae, the only species in the genus Anasis. [more]

Anatrichis

[more]

Anatrichus

[more]

Anchista

Anchista is a genus of beetles in the family Carabidae, containing the following species: [more]

Anchomenidius

[more]

Anchomenus

Anchomenus is a genus of ground beetle native to the Palearctic (including Europe), the Near East and North Africa. [more]

Anchonoderus

Anchonoderus is a genus of beetles in the family Carabidae, containing the following species: [more]

Anchonymus

[more]

Anchotefflus

[more]

Anchus

[more]

Anillaspis

[more]

Anillinus

[more]

Anillodes

[more]

Anillus

[more]

Anisodactylus

Anisodactylus is a genus of ground beetle native to the Palearctic (including Europe), the Near East and North Africa. It contains the following species: [more]

Anomoglossus

[more]

Anomotarus

[more]

Anophthalmus

Anophthalmus is a genus of ground beetle endemic to Europe. [more]

Anoplogenius

[more]

Anthia

Anthia is a genus of the ground beetle family, the Carabidae. Species of Anthia can spray a jet of formic acid up to 30 centimetres (12 in), which if not treated, can cause blindness in cats and chickens. [more]

Anthracus

Anthracus is a genus of beetles in the family Carabidae, containing the following species: [more]

Antisphodrus

[more]

Apatrobus

[more]

Apenes

Apenes is a genus of beetles in the family Carabidae, containing the following species: [more]

Aphaenopidius

[more]

Aphaenops

[more]

Aphaenotyphlus

[more]

Aphanotrechus

[more]

Aphoenops

[more]

Apoduvalius

[more]

Apotomopterus

[more]

Apristus

Apristus is a genus of beetles in the family Carabidae, containing the following species: [more]

Apteromesus

[more]

Aptinus

Aptinus is a genus of ground beetle native to Europe and the Near East. It contains the following species: [more]

Archicolliuris

[more]

Archipatrobus

[more]

Arctaphaenops

[more]

Ardistomis

[more]

Argutor

[more]

Armatocillenus

[more]

Arthropterus

Arthropterus is a genus of beetles in the family Carabidae, containing the following species: [more]

Asaphidion

Asaphidion is a Holarctic genus of ground beetle native to Europe, the Near East, North Africa, and North America. [more]

Aspidoglossa

[more]

Atelothrus

[more]

Atrachycnemis

[more]

Atractonotus

Atractonotus mulsanti is a species of beetle in the family Carabidae, the only species in the genus Atractonotus. [more]

Atranus

[more]

Aulacoryssus

[more]

Axinidium

[more]

Axinopalpus

Axinopalpus is a genus of beetles in the family Carabidae, containing the following species: [more]

Badister

Badister is a genus of ground beetle in the Carabidae family native to North Africa, the Near East and the Palearctic, including Europe. [more]

Baeoglossa

[more]

Baryneus

[more]

Barypristus

[more]

Barysomus

[more]

Basistichus

[more]

Batenus

[more]

Baudia

[more]

Bembidion

[more]

Binaghites

[more]

Blackburnia

[more]

Blemus

[more]

Blethisa

[more]

Boldoriella

[more]

Bothriopterus

[more]

Bothynotrechus

[more]

Brachichila

[more]

Brachinus

[more]

Bradycellus

[more]

Brasiella

[more]

Broscodera

[more]

Brosconymus

[more]

Broscosoma

[more]

Broscus

[more]

Caelostomus

[more]

Cainogenion

[more]

Calathidius

[more]

Calathus

A Genus in the Kingdom Animalia.[1] [more]

Calleida

[more]

Callida

[more]

Callisthenes

A Genus in the Kingdom Animalia.[2] [more]

Callistoides

[more]

Callistomimus

[more]

Callistus

Callistus, Callixtus (both Latin) and Kallistos (in Greek) may refer to: [more]

Callytron

[more]

Calodromius

[more]

Calomera

[more]

Calophaena

[more]

Calosama

[more]

Calosoma

[more]

Calybe

[more]

Campalita

[more]

Canarobius

[more]

Carabites

[more]

Carabomorphus

[more]

Carabus

Carabus is a genus of beetle in family Carabidae. [more]

Cardiaderus

[more]

Cardiomera

[more]

Carenum

[more]

Cassolaia

[more]

Castelnaudia

[more]

Catadromus

[more]

Catascopus

[more]

Cedrorum

[more]

Celaenephes

[more]

Cephalota

[more]

Ceroglossus

[more]

Cerotalis

[more]

Chaetocarabus

[more]

Chaetodera

[more]

Chalcomenus

[more]

Cheporus

[more]

Chinajujiroa

[more]

Chlaeniellus

[more]

Chlaeniomimus

[more]

Chlaenius

[more]

Chrysocarabus

[more]

Chylnus

[more]

Cicindela

Cicindela are generally brightly colored and metallic beetles, often with some sort of patterning of ivory or cream-colored markings. They are most abundant and diverse in habitats with sandy soil (though some prefer clay), and very often near bodies of water, even if seasonally transient; along river, sea and lake shores, on sand dunes, around playa lakebeds and on clay banks or woodland paths. [more]

Cicindella

[more]

Cillenus

[more]

Clarencia

[more]

Clivina

[more]

Clivinia

[more]

Colfax

[more]

Colliuris

[more]

Colpocaccus

[more]

Colpodes

[more]

Colpodiscus

[more]

Conopterum

[more]

Coptocarpus

Coptodera

[more]

Coptodere

Coptolabrus

[more]

Corsyra

Cosmodela

[more]

Cosmodiscus

Crasodactylus

Craspedonotus

Craspedophorus

Cratacanthus

[more]

Cratoferonia

Cratogaster

Croatotrechus

Cryptophonus

Ctenosta

[more]

Ctenostoma

Cuneipectus

[more]

Curtonotus

[more]

Cychrus

[more]

Cyclotrachelus

[more]

Cylindera

[more]

Cylindronotum

[more]

Cymbionotum

Cymindis

[more]

Cypholoba

[more]

Dalmataphaenops

Dalyat

Damaster

[more]

Daptus

[more]

Darlingtonea

[more]

Darodilia

Decogmus

Delichoctis

Delinius

Deltomerus

[more]

Demetrias

A Genus in the Kingdom Animalia. [more]

Demetrida

[more]

Dercylinus

[more]

Derobroscus

[more]

Deropristus

[more]

Desera

[more]

Dhanya

[more]

Diacheila

[more]

Diachromus

[more]

Dicaelus

[more]

Dicheirotrichus

[more]

Dicheirus

[more]

Dichrochile

Dicranoglossus

Dicranoncus

[more]

Dicranonus

Dicraspeda

[more]

Dicrodontus

Dineutes

[more]

Dinodes

Diplochaetus

[more]

Diplocheila

[more]

Diplochelia

Diplous

[more]

Disamara

[more]

Dischissus

Discoderus

[more]

Disenochus

[more]

Distichus

Distipsidera

Ditomus

[more]

Dixus

Doderotrechus

Dolichoctis

[more]

Dolichus

[more]

Dolischus

Drimostoma

Dromochorus

[more]

Drypta

[more]

Duvaliopsis

[more]

Duvalius

[more]

Dyschiriodes

[more]

Dyschirius

[more]

Dyschirus

[more]

Dystrichothorax

Eccoptomenus

[more]

Eccoptoptera

[more]

Ega

A Genus in the Kingdom Animalia.[3] [more]

Egadroma

[more]

Elaphropus

[more]

Elaphrus

[more]

Emphanes

[more]

Enceladus

[more]

Endynomena

[more]

Enkelgezum

Entomoantyx

Eobroscus

Eocarterus

Eochlaenius

Epaphiopsis

Epaphius

[more]

Epilectus

Episcopellus

[more]

Epomis

Eripus

[more]

Eucaerus

[more]

Eucarterus

Eucolliuris

Eudalia

[more]

Eugrapha

[more]

Euphorticus

[more]

Euplynes

Euproctinus

[more]

Europhilus

[more]

Euryaptus

Eurycarabus

Euryderus

[more]

Eurylychnus

[more]

Eurylynchus

Eurynebria

Euryscaphus

Eurysoma

Eurystomis

Euschizomerus

Eusthulmannium

Euthenarus

Eutoma

Eutrechopsis

Eutrechus

Eutrichopus

[more]

Evolenes

[more]

Formosiella

Fortagonum

[more]

Galaicodytes

Galerita

[more]

Galeritula

[more]

Galiciotyphlotes

Gastrellarius

[more]

Gatrellarius

Gehringia

[more]

Geobaenus

[more]

Geopinus

[more]

Geoscaptus

[more]

Gigadema

[more]

Glabrotrechus

Gnathaphanus

[more]

Gnatholymnaeum

[more]

Gnathoxys

Goniotropus

[more]

Gonogenia

[more]

Graniger

Graphipterus

Gynandromorphus

Gynandropus

[more]

Habrodera

[more]

Hadrocarabus

[more]

Halocoryza

[more]

Haolochlaenius

Haplaner

Haplochiaenius

Haplochlaenius

Harpalellus

[more]

Harpalobrachys

[more]

Harpalus

A Genus in the Kingdom Animalia.[4] [more]

Harplaner

Hartonymus

[more]

Helluapterus

Helluarchus

Helluodema

Helluomorphoides

[more]

Helluonidius

Hemicarabus

Henrotius

Hiletus

Hispalis

Holcoderus

[more]

Hololeius

Homethes

Homodela

[more]

Hormacrus

Horologion

A Genus in the Kingdom Animalia. [more]

Hydraphaenops

[more]

Hydrotrechus

Hygrocarabus

Hyparpalus

[more]

Hyperion

Hypharpax

[more]

Hypotyphlus

Iberanillus

Iberodytes

Iberotrechus

Idacarabus

Iga

Ildobates

Illaphanus

Infernophilus

[more]

Inna

[more]

Ishidatrechus

Italaphaenops

Italodytes

Jdiochroma

Jujiroa

[more]

Laccocenus

Laccopterum

Laccoscaphus

Lachnocrepis

[more]

Lachnoderma

Lachnolebia

Lachnophorus

[more]

Lacordairia

Laemostenus

[more]

Lamprias

Lasiotrechus

[more]

Lauosterus

Lebia

[more]

Lebidia

Lecalida

[more]

Lecanomerus

[more]

Leiradira

Leistus

[more]

Leptocarabus

Lessinodytes

Lesticus

[more]

Lestignathus

Licinopsis

[more]

Licinus

[more]

Limnastis

[more]

Limodromus

Lindrothius

[more]

Lionychus

[more]

Lioptera

Lipaster

Lissopogonus

Lithochlaenius

Lonchosternus

Lophoglossus

[more]

Lophyra

[more]

Lophyridia

[more]

Loricera

[more]

Lorocera

Lorostema

Lovricia

Loxandrus

[more]

Loxodactylus

Loxogenius

Loxoncus

Luperca

[more]

Lymnaeum

Lymnastis

Lymnastus

[more]

Lymneops

[more]

Mantichora

[more]

Masoreus

[more]

Mauna

[more]

Mecodema

Mecodema is a genus of ground beetle in the family Carabidae. [more]

Mecomenus

[more]

Mecyclothorax

[more]

Mecynognathus

Megacephala

[more]

Megalopaussus

Melisodera

Meonis

[more]

Mesolestes

Mesothriscus

[more]

Metabletus

[more]

Metacolpodes

Metadromius

Metallina

[more]

Metrius

[more]

Metromenus

[more]

Micralymma

A Genus in the Kingdom Animalia. [more]

Micratopus

[more]

Micrixys

[more]

Microcosmodes

Microderes

Microferonia

Microleser

Microlestes

[more]

Microlestodes

Microtyphlus

Migadops

[more]

Mimotrechus

Minypatrobus

Mioptachys

[more]

Miscelus

[more]

Miscodera

[more]

Molopidius

Molops

[more]

Monocentrum

Morimotoidius

Moriodema

Morion

A Genus in the Kingdom Animalia.[5] [more]

Moriosomus

Mormolyce

[more]

Mouhotia

Muscelus

Myas

[more]

Myriochila

Myriochile

[more]

Myrmecodemus

Mysticomenus

[more]

Mystropomus

Nabria

Neaphaenops

[more]

Nebria

[more]

Nelsonites

[more]

Nemotarsus

[more]

Neocarenum

Neoeudalia

Neohelluo

Neolaphyra

[more]

Neonomius

Neoscaphus

Neotrechus

Nesacinopus

Nesorthomus

Netrodera

[more]

Nickerlea

Nippononebria

Nomius

Apollo (Attic, Ionic, and Homeric Greek: , Apollon (gen.: ?p???????); Doric: ?p?????, Apellon; Arcadocypriot: ?pe????, Apeilon; Aeolic: ?p????, Aploun; Latin: Apollo) is one of the most important and complex of the Olympian deities in ancient Greek and Roman religion, Greco?Roman Neopaganism, and Greek and Roman mythology. The ideal of the kouros (a beardless, athletic youth), Apollo has been variously recognized as a god of light and the sun, truth and prophecy, healing, plague, music, poetry, and more. Apollo is the son of Zeus and Leto, and has a twin sister, the chaste huntress Artemis. Apollo is known in Greek-influenced Etruscan mythology as Apulu. [more]

Notagonum

[more]

Notaphus

[more]

Nothiophilus

Notiobia

[more]

Notiophilus

[more]

Notonomus

[more]

Notoplatynus

Notospeophonus

Nototarsus

Nototrechus

Nurus

Ocyropus

Ocys

[more]

Odacantha

[more]

Odacanthella

Odacentha

Odontium

[more]

Odontocarus

[more]

Odontocheila

[more]

Odontonyx

[more]

Oedesis

Ohomopterus

[more]

Oliskhopus

Olisthopus

[more]

Omophron

[more]

Omphra

[more]

Omphreus

Omus

Omus is a genus of , subfamily Cicindelinae. Its members are dark colored, nocturnal, and flightless. All members occur along the west coast of North America. [more]

Onota

Onycholabis

Onypterygia

Oodes

[more]

Ophionea

Ophonus

[more]

Opisthius

[more]

Oreonebria

Orinicarabus

Orinocarabus

Orionella

Orotrechus

[more]

Orthogonius

Orthomus

[more]

Orzolina

Oscadytes

Oxycentrus

Oxycheila

Oxycrepis

[more]

Oxydrepanus

Oxyglychus

Oxygonia

Oxypselaphus

[more]

Ozaena

A Genus in the Kingdom Animalia. [more]

Pachydinodes

[more]

Pachyteles

[more]

Pamborus

Panagaeus

[more]

Pangus

Parabroscus

Paradromius

[more]

Paraeutrichopus

Parallelomorphus

Paranchus

[more]

Paraphaenops

Paratachys

[more]

Parazuphium

[more]

Parena

[more]

Parophonus

[more]

Parroa

Pasimachus

[more]

Passalidius

[more]

Patrobus

[more]

Paussus

Pediomorphus

Pedius

Pelmatellus

[more]

Pelophila

[more]

Penetretus

Pentacomia

Pentagonica

[more]

Percosia

[more]

Percosoma

Percus

Pericalus

[more]

Pericompsus

[more]

Pericompus

[more]

Perigona

[more]

Perileptus

[more]

Peripristus

Peripristus ater is a species of beetle in the family Carabidae, the only species in the genus Peripristus. [more]

Peronomerus

Pheggomisetes

Pheropsophus

[more]

Philochthus

[more]

Philorhizus

[more]

Philoscaphus

Phloeoxena

[more]

Phloeozeteus

Phorticosomus

Phrypeus

[more]

Physea

Piesmus

[more]

Pinacodera

[more]

Piosoma

Piosoma setosum is a species of beetle in the family Carabidae, the only species in the genus Piosoma. [more]

Plagiotelum

Plagiotelum is a genus of beetles in the family Carabidae, containing the following species: [more]

Plataphus

Platidiolus

[more]

Platus

[more]

Platycarabus

Platycerozaena

[more]

Platyderus

[more]

Platylytron

Platynus

[more]

Platypatrobus

[more]

Platysma

A Genus in the Kingdom Animalia. [more]

Platytarus

Plochinus

Plochionus

[more]

Poecilus

[more]

Pogonistes

[more]

Pogonodaptus

[more]

Pogonoglossus

Pogonus

[more]

Poliocypas

Polistichus

Polpochila

[more]

Polyderis

[more]

Polystichus

[more]

Porotachys

[more]

Prena

Princidium

Pristonychus

[more]

Pristosia

Procarabus

Procerus

A Genus in the Kingdom Animalia.[6] [more]

Procrustes

A Genus in the Kingdom Animalia. [more]

Procustes

Prodisenochus

[more]

Promecoderus

Promecognathus

[more]

Prosopogmus

[more]

Pseudabarys

[more]

Pseudanophthalmus

[more]

Pseudaptinus

[more]

Pseudobroscus

[more]

Pseudomasoreus

Pseudomorpha

[more]

Pseudomyas

Pseudoophonus

Pseudoopterus

[more]

Pseudotaphoxenus

Pseudotrechus

Pseudoxycheila

Psydrus

[more]

Pterostichus

[more]

Quammenis

Rambousekiella

Reicheia

Reicheiodes

Rembus

[more]

Rhadine

[more]

Rhegmatobius

Rhopalostyla

Rhytisternus

Risophilus

Rupa

Sakagutia

Sardaphaenops

[more]

Sarothrocrepis

Sarticus

Scambocarabus

Scaphinotus

[more]

Scaraphites

[more]

Scarites

[more]

Schizogenius

[more]

Scopodes

[more]

Scotodipnus

Scotoplanetes

Scybalicus

Selenophorus

[more]

Selina

Semiardistomis

Semiophonus

Sericoda

Sericoda is a of ground beetles. They are pyrophilous, meaning that they are attracted by burned areas of forest, where they are found in large numbers the summer following the fires, after which their numbers decrease. [more]

Simodontus

Sinechostictus

Singilis

Sinocaralosoma

Sirdenus

Sloaneana

Somatrichus

[more]

Somoplatus

Somotrichus

[more]

Spelaeodytes

Spelaeovulcania

Speleotyphlus

Speluncarius

Speomolops

Speotrechus

Sphaeroderus

[more]

Sphaerotachys

Sphallomorpha

[more]

Spheroderus

Sphodropsis

Sphodrus

[more]

Stenocrepis

[more]

Stenolophus

[more]

Stenomorphus

[more]

Stereocerus

[more]

Stichonotus

[more]

Stolonis

[more]

Stomis

[more]

Stricklandia

[more]

Stylulus

[more]

Styracoderus

Synchus

[more]

Syntomus

[more]

Synuchidius

Synuchus

[more]

Tachys

[more]

Tachyta

[more]

Tachyura

[more]

Taicona

Tanystoma

Tanythrix

Taphoxenus

Tapinopterus

Tasmanitachoides

Tecnophilus

[more]

Tefflus

[more]

Tetracha

[more]

Tetragonoderus

[more]

Tetraleucus

[more]

Thalassophilus

[more]

Thalassotrechus

[more]

Thalpius

22503 Thalpius (1997 TB12) is a Jupiter Trojan discovered on October 7, 1997 by M. Tichy and Z. Moravec at Klet. [more]

Therates

[more]

Thermophilum

Anthia is a genus of the ground beetle family, the Carabidae. Species of Anthia can spray a jet of formic acid up to 30 centimetres (12 in), which if not treated, can cause blindness in cats and chickens. [more]

Thermoscelis

[more]

Thopeutica

Thopeutica is a genus of beetles in the family Carabidae, containing the following species: [more]

Thyreopterus

Thyreopterus is a genus of beetles in the family Carabidae, containing the following species: [more]

Tienmutrechus

Tinautius

Tinoderus

Tomocarabus

Trechiama

Trechiana

[more]

Trechicus

Trechimorphus

Trechoblemus

[more]

Trechus

[more]

Trepanes

Trephionus

Trichaphaenops

[more]

Trichis

Trichocellus

[more]

Trichosternus

Trichotichnus

[more]

Tricondyla

[more]

Trigonognatha

Trigonotoma

Tritrichis

Troglanillus

[more]

Troglorites

Trymosternus

Tschitscherinellus

Typhlocharis

Typhlochoromus

Typhlonesiotes

[more]

Typhloreicheia

Typhlotrechus

Typsiharpalus

Vachinius

Wolltinerfia

[more]

Xenion

[more]

Xenotrechus

[more]

Xestonotus

[more]

Zabrus

[more]

Zacotus

[more]

Zargus

Zariquieya

Zecillenus

[more]

Zeodera

Zuphium

[more]

At least 14 species and subspecies belong to the Genus Zuphium.

More info about the Genus Zuphium may be found here.

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. ^ Shozo Osawa, Zhi-Hui Su & Yuki Inmura (2004). Molecular Phylogeny and Evolution of Carabid Ground Beetles. Springer. ISBN 4-431-00487-4. 
  9. ^ I. L?bl & ; 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 bee tles 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. 

Footnotes

  1. http://bugguide.net/index.php?q=search&keys=Calathus&search=Search
  2. http://bugguide.net/index.php?q=search&keys=Callisthenes&search=Search
  3. http://bugguide.net/index.php?q=search&keys=Ega&search=Search
  4. http://bugguide.net/index.php?q=search&keys=Harpalus&search=Search
  5. http://bugguide.net/index.php?q=search&keys=Morion&search=Search
  6. http://bugguide.net/index.php?q=search&keys=Procerus&search=Search
  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]

Further Reading

  • E. Csiki (1946) (in German). Die K?ferfauna des Karpaten-Beckens [The beetle fauna of the Carparthian basin]. Budapest. pp. 71?546. 
  • K. Kult (1947) (in Czech). Kl?c k urcov?n? brouku celedi Carabidae Ceskoslovensk? 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 K?fer des Deutschen Reiches [The beetles of the German Empire]. Stuttgart: K. G. Lutz. 

External links

Sources

  • The text on this page is licensed under the GNU Free Documentation License. It includes material from Wikipedia retrieved Wednesday, April 25, 2012.
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Last Revised: August 24, 2012
2012/08/24 13:26:51