Overview
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]
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]
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.
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
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
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?
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]
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.
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
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
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
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
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.
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
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
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?
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]
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.
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
Carabinae La treille, 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
Omophronin ae 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
Amblytelinae Sloane, 1898[Note 1]
- Amblytelus< /li>
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?
- Gehringiini ? Trechinae or a distinct subfamily
- Metiini ? Harpalinae?
Notes
References
- ^ "Carabidae Taxa". Carabidae of the World. 2011. http://carabidae.pro/carabidae. Retrieved 24 Jun 2011.
- ^ 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.
- ^ John L. Capinera. Encyclopedia of Entomology. p. 1746.
- ^ 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.
- ^ "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.
- ^ 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.
- ^ Charles Darwin (1846). "Lett er to Leonard Jenyns, October 17, 1846". http://www.darwinproject.ac.uk/darwinletters/calendar/entry-1009.html.
- ^ Shozo Osawa, Zhi-Hui Su & Yuki Inmura (2004). Molecular Phylogeny and Evolution of Carabid Ground Beetles. Springer. ISBN 4-431-00487-4.
- ^ I. L?bl & A. Smetana, ed. (2003?). Catalogue of Palaearctic Coleoptera. Stenstrup, Denmark: Apollo Books.
- ^ "Trees of family Carabidae". Carabidae of the World Database. 2008. http://www.carabidae.ru/carabidae.htm. Retrieved July 24, 2008.
- ^ 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.
- ^ 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.
- ^ "Harpalinae". Fauna Europaea. 2004. http://www.faunaeur.org/full_res ults.php?id=379457. Retrieved February 16, 2011.
- ^ 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:
- Subfamily (43): Aephnidiinae · Agoninae · Amblystominae · Anchonoderinae · Anisodactylinae · Badistrinae · Bembidiinae · Brachininae · Broscinae · Callidinae · Callistinae · Carabinae · Cicindelinae · Colliurinae · Cychrinae · Cymindinae · Demetriinae · Ditominae · Dromiinae · Dryptinae · Elaphrinae · Harpalinae · Lebiinae · Licininae · Loricerinae · Mormolycinae · Nebriinae · Notiophilinae · Octopodinae · Odacanthinae · Omophroninae · Oodinae · Panagaeinae · P atrobinae · Pentagonicinae · Perigoninae · Pogoninae · Pterostichinae · Scaritinae · Stenolophinae · Trechinae · Zabrinae · Zuphiinae
- Tribe (6): Bembidiini · Helluonini · Limnastini · Panagaeini · Pterostichini · Tachyini
- Genus (765): Abacetus · Abacidus · Abax · Abroscelis · Acanthoscelis · Acinopus · Acupalpus · Adelotopus · Adriaphaenops · Aephnidius · Aepopsis · Aepus · Agatus · Agelaea · Agonocheila · Agonoderus · Agonotrechus · Agonum · Agostenus · Agra · Ainocolpodes · Albanotrechus · Allegrettia · Allotriopus · Altagonum · Amara · Amaroschema · Amblycheila · Amblygnathus · Amblystomus · Amblytelus · Ambylgnathus · Amerinus · Amerizus · Ameroduvalius · Amphasia · Amphimenes · Anasis · Anatrichis · Anatrichus · Anchista · Anchomenidius · Anchomenus · Anchonoderus · Anchonymus · Anchotefflus · Anchus · Anillaspis · Anillinus · Anillodes · Anillus · Anisodactylus · Anomoglossus · Anomotarus · Anophthalmus · Anoplogenius · Anthia · Anthracus · Antisphodrus · Apatrobus · Apenes · Aphaenopidius · Aphaenops · Aphaenotyphlus · Aphanotrechus · Aphoenops · Apoduvalius · Apotomopterus · Apristus · Apteromesus · Aptinus · Archicolliuris · Archipatrobus · Arctaphaenops · Ardistomis · Argutor · Armatocillenus · Arthropterus · Asaphidion · Aspidoglossa · Atelothrus · Atrachycnemis · Atractonotus · Atranus · Aulacoryssus · Axinidium · Axinopalpus · Badister · Baeoglossa · Baryneus · Barypristus · Barysomus · Basistichus · Batenus · Baudia · Bembidion · Binaghites · Blackburnia · Blemus · Blethisa · Boldoriella · Bothriopterus · Bothynotrechus · Brachichila · Brachinus · Bradycellus · Brasiella · Broscodera · Brosconymus · Broscosoma · Broscus · Caelostomus · Cainogenion · Calathidius · Calathus · Calleida · Callida · Callisthenes · Callistoides · Callistomimus · Callistus · Callytron · Calodromius · Calomera · Calophaena · Calosama · Calosoma · Calybe · Campalita · Canarobius · Carabites · Carabomorphus · Carabus · Cardiaderus · Cardiomera · Carenum · Cassolaia · Castelnaudia · Catadromus · Catascopus · Cedrorum · Celaenephes · Cephalota · Ceroglossus · Cerotalis · Chaetocarabus · Chaetodera · Chalcomenus · Cheporus · Chinajujiroa · Chlaeniellus · Chlaeniomimus · Chlaenius · Chrysocarabus · Chylnus · Cicindela · Cicindella · Cillenus · Clarencia · Clivina · Clivinia · Colfax · Colliuris · Colpocaccus · Colpodes · Colpodiscus · Conopterum · Coptocarpus · Coptodera · Coptodere · Coptolabrus · Corsyra · Cosmodela · Cosmodiscus · Crasodactylus · Craspedonotus · Craspedophorus · Cratacanthus · Cratoferonia · Cratogaster · Croatotrechus · Cryptophonus · Ctenosta · Ctenostoma · Cuneipectus · Curtonotus · Cychrus · Cyclotrachelus · Cylindera · Cylindronotum · Cymbionotum · Cymindis · Cypholoba · Dalmataphaenops · Dalyat · Damaster · Daptus · Darlingtonea · Darodilia · Decogmus · Delichoctis · Delinius · Deltomerus · Demetrias · Demetrida · Dercylinus · Derobroscus · Deropristus · Desera · Dhanya · Diacheila · Diachromus · Dicaelus · Dicheirotrichus · Dicheirus · Dichrochile · Dicranoglossus · Dicranoncus · Dicranonus · Dicraspeda · Dicrodontus · Dineutes · Dinodes · Diplochaetus · Diplocheila · Diplochelia · Diplous · Disamara · Dischissus · Discoderus · Disenochus · Distichus · Distipsidera · Ditomus · Dixus · Doderotrechus · Dolichoctis · Dolichus · Dolischus · Drimostoma · Dromochorus · Drypta · Duvaliopsis · Duvalius · Dyschiriodes · Dyschirius · Dyschirus · Dystrichothorax · Eccoptomenus · Eccoptoptera · Ega · Egadroma · Elaphropus · Elaphrus · Emphanes · Enceladus · Endynomena · Enkelgezum · Entomoantyx · Eobroscus · Eocarterus · Eochlaenius · Epaphiopsis · Epaphius · Epilectus · Episcopellus · Epomis · Eripus · Eucaerus · Eucarterus · Eucolliuris · Eudalia · Eugrapha · Euphorticus · Euplynes · Euproctinus · Europhilus · Euryaptus · Eurycarabus · Euryderus · Eurylychnus · Eurylynchus · Eurynebria · Euryscaphus · Eurysoma · Eurystomis · Euschizomerus · Eusthulmannium · Euthenarus · Eutoma · Eutrechopsis · Eutrechus · Eutrichopus · Evolenes · Formosiella · Fortagonum · Galaicodytes · Galerita · Galeritula · Galiciotyphlotes · Gastrellarius · Gatrellarius · Gehringia · Geobaenus · Geopinus · Geoscaptus · Gigadema · Glabrotrechus · Gnathaphanus · Gnatholymnaeum · Gnathoxys · Goniotropus · Gonogenia · Graniger · Graphipterus · Gynandromorphus · Gynandropus · Habrodera · Hadrocarabus · Halocoryza · Haolochlaenius · Haplaner · Haplochiaenius · Haplochlaenius · Harpalellus · Harpalobrachys · Harpalus · Harplaner · Hartonymus · Helluapterus · Helluarchus · Helluodema · Helluomorphoides · Helluonidius · Hemicarabus · Henrotius · Hiletus · Hispalis · Holcoderus · Hololeius · Homethes · Homodela · Hormacrus · Horologion · Hydraphaenops · Hydrotrechus · Hygrocarabus · Hyparpalus · Hyperion · Hypharpax · Hypotyphlus · Iberanillus · Iberodytes · Iberotrechus · Idacarabus · Iga · Ildobates · Illaphanus · Infernophilus · Inna · Ishidatrechus · Italaphaenops · Italodytes · Jdiochroma · Jujiroa · Laccocenus · Laccopterum · Laccoscaphus · Lachnocrepis · Lachnoderma · Lachnolebia · Lachnophorus · Lacordairia · Laemostenus · Lamprias · Lasiotrechus · Lauosterus · Lebia · Lebidia · Lecalida · Lecanomerus · Leiradira · Leistus · Leptocarabus · Lessinodytes · Lesticus · Lestignathus · Licinopsis · Licinus · Limnastis · Limodromus · Lindrothius · Lionychus · Lioptera · Lipaster · Lissopogonus · Lithochlaenius · Lonchosternus · Lophoglossus · Lophyra · Lophyridia · Loricera · Lorocera · Lorostema · Lovricia · Loxandrus · Loxodactylus · Loxogenius · Loxoncus · Luperca · Lymnaeum · Lymnastis · Lymnastus · Lymneops · Mantichora · Masoreus · Mauna · Mecodema · Mecomenus · Mecyclothorax · Mecynognathus · Megacephala · Megalopaussus · Melisodera · Meonis · Mesolestes · Mesothriscus · Metabletus · Metacolpodes · Metadromius · Metallina · Metrius · Metromenus · Micralymma · Micratopus · Micrixys · Microcosmodes · Microderes · Microferonia · Microleser · Microlestes · Microlestodes · Microtyphlus · Migadops · Mimotrechus · Minypatrobus · Mioptachys · Miscelus · Miscodera · Molopidius · Molops · Monocentrum · Morimotoidius · Moriodema · Morion · Moriosomus · Mormolyce · Mouhotia · Muscelus · Myas · Myriochila · Myriochile · Myrmecodemus · Mysticomenus · Mystropomus · Nabria · Neaphaenops · Nebria · Nelsonites · Nemotarsus · Neocarenum · Neoeudalia · Neohelluo · Neolaphyra · Neonomius · Neoscaphus · Neotrechus · Nesacinopus · Nesorthomus · Netrodera · Nickerlea · Nippononebria · Nomius · Notagonum · Notaphus · Nothiophilus · Notiobia · Notiophilus · Notonomus · Notoplatynus · Notospeophonus · Nototarsus · Nototrechus · Nurus · Ocyropus · Ocys · Odacantha · Odacanthella · Odacentha · Odontium · Odontocarus · Odontocheila · Odontonyx · Oedesis · Ohomopterus · Oliskhopus · Olisthopus · Omophron · Omphra · Omphreus · Omus · Onota · Onycholabis · Onypterygia · Oodes · Ophionea · Ophonus · Opisthius · Oreonebria · Orinicarabus · Orinocarabus · Orionella · Orotrechus · Orthogonius · Orthomus · Orzolina · Oscadytes · Oxycentrus · Oxycheila · Oxycrepis · Oxydrepanus · Oxyglychus · Oxygonia · Oxypselaphus · Ozaena · Pachydinodes · Pachyteles · Pamborus · Panagaeus · Pangus · Parabroscus · Paradromius · Paraeutrichopus · Parallelomorphus · Paranchus · Paraphaenops · Paratachys · Parazuphium · Parena · Parophonus · Parroa · Pasimachus · Passalidius · Patrobus · Paussus · Pediomorphus · Pedius · Pelmatellus · Pelophila · Penetretus · Pentacomia · Pentagonica · Percosia · Percosoma · Percus · Pericalus · Pericompsus · Pericompus · Perigona · Perileptus · Peripristus · Peronomerus · Pheggomisetes · Pheropsophus · Philochthus · Philorhizus · Philoscaphus · Phloeoxena · Phloeozeteus · Phorticosomus · Phrypeus · Physea · Piesmus · Pinacodera · Piosoma · Plagiotelum · Plataphus · Platidiolus · Platus · Platycarabus · Platycerozaena · Platyderus · Platylytron · Platynus · Platypatrobus · Platysma · Platytarus · Plochinus · Plochionus · Poecilus · Pogonistes · Pogonodaptus · Pogonoglossus · Pogonus · Poliocypas · Polistichus · Polpochila · Polyderis · Polystichus · Porotachys · Prena · Princidium · Pristonychus · Pristosia · Procarabus · Procerus · Procrustes · Procustes · Prodisenochus · Promecoderus · Promecognathus · Prosopogmus · Pseudabarys · Pseudanophthalmus · Pseudaptinus · Pseudobroscus · Pseudomasoreus · Pseudomorpha · Pseudomyas · Pseudoophonus · Pseudoopterus · Pseudotaphoxenus · Pseudotrechus · Pseudoxycheila · Psydrus · Pterostichus · Quammenis · Rambousekiella · Reicheia · Reicheiodes · Rembus · Rhadine · Rhegmatobius · Rhopalostyla · Rhytisternus · Risophilus · Rupa · Sakagutia · Sardaphaenops · Sarothrocrepis · Sarticus · Scambocarabus · Scaphinotus · Scaraphites · Scarites · Schizogenius · Scopodes · Scotodipnus · Scotoplanetes · Scybalicus · Selenophorus · Selina · Semiardistomis · Semiophonus · Sericoda · Simodontus · Sinechostictus · Singilis · Sinocaralosoma · Sirdenus · Sloaneana · Somatrichus · Somoplatus · Somotrichus · Spelaeodytes · Spelaeovulcania · Speleotyphlus · Speluncarius · Speomolops · Speotrechus · Sphaeroderus · Sphaerotachys · Sphallomorpha · Spheroderus · Sphodropsis · Sphodrus · Stenocrepis · Stenolophus · Stenomorphus · Stereocerus · Stichonotus · Stolonis · Stomis · Stricklandia · Stylulus · Styracoderus · Synchus · Syntomus · Synuchidius · Synuchus · Tachys · Tachyta · Tachyura · Taicona · Tanystoma · Tanythrix · Taphoxenus · Tapinopterus · Tasmanitachoides · Tecnophilus · Tefflus · Tetracha · Tetragonoderus · Tetraleucus · Thalassophilus · Thalassotrechus · Thalpius · Therates · Thermophilum · Thermoscelis · Thopeutica · Thyreopterus · Tienmutrechus · Tinautius · Tinoderus · Tomocarabus · Trechiama · Trechiana · Trechicus · Trechimorphus · Trechoblemus · Trechus · Trepanes · Trephionus · Trichaphaenops · Trichis · Trichocellus · Trichosternus · Trichotichnus · Tricondyla · Trigonognatha · Trigonotoma · Tritrichis · Troglanillus · Troglorites · Trymosternus · Tschitscherinellus · Typhlocharis · Typhlochoromus · Typhlonesiotes · Typhloreicheia · Typhlotrechus · Typsiharpalus · Vachinius · Wolltinerfia · Xenion · Xenotrechus · Xestonotus · Zabrus · Zacotus · Zargus · Zariquieya · Zecillenus · Zeodera · Zuphium
- Species: ZipcodeZoo has pages for 10,257 species and subspecies in the Family Carabidae.
Genera
Abacetus
Abacidus
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
Acinopus
Acupalpus
Acupalpus is a genus of insect-eating beetle species. Its representatives are found across Europe and Asia. [more]
Adelotopus
Adriaphaenops
Aephnidius
Aepopsis
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
Agelaea
Agelaea is a genus of beetles in the family Carabidae, containing the following species: [more]
Agonocheila
Agonoderus
Agonotrechus
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
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
Albanotrechus
Allegrettia
Allotriopus
Altagonum
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
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
Amblystomus
Amblystomus is a genus of beetles in the family Carabidae, containing the following species: [more]
Amblytelus
Ambylgnathus
Amerinus
Amerinus linearis is a species of beetle in the family Carabidae, the only species in the genus Amerinus. [more]
Amerizus
Ameroduvalius
Amphasia
Amphasia is a genus of beetles in the family Carabidae, containing the following species: [more]
Amphimenes
Anasis
Anasis howitti is a species of beetle in the family Carabidae, the only species in the genus Anasis. [more]
Anatrichis
Anatrichus
Anchista
Anchista is a genus of beetles in the family Carabidae, containing the following species: [more]
Anchomenidius
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
Anchotefflus
Anchus
Anillaspis
Anillinus
Anillodes
Anillus
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
Anomotarus
Anophthalmus
Anophthalmus is a genus of ground beetle endemic to Europe. [more]
Anoplogenius
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
Apatrobus
Apenes
Apenes is a genus of beetles in the family Carabidae, containing the following species: [more]
Aphaenopidius
Aphaenops
Aphaenotyphlus
Aphanotrechus
Aphoenops
Apoduvalius
Apotomopterus
Apristus
Apristus is a genus of beetles in the family Carabidae, containing the following species: [more]
Apteromesus
Aptinus
Aptinus is a genus of ground beetle native to Europe and the Near East. It contains the following species: [more]
Archicolliuris
Archipatrobus
Arctaphaenops
Ardistomis
Argutor
Armatocillenus
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
Atelothrus
Atrachycnemis
Atractonotus
Atractonotus mulsanti is a species of beetle in the family Carabidae, the only species in the genus Atractonotus. [more]
Atranus
Aulacoryssus
Axinidium
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
Baryneus
Barypristus
Barysomus
Basistichus
Batenus
Baudia
Bembidion
Binaghites
Blackburnia
Blemus
Blethisa
Boldoriella
Bothriopterus
Bothynotrechus
Brachichila
Brachinus
Bradycellus
Brasiella
Broscodera
Brosconymus
Broscosoma
Broscus
Caelostomus
Cainogenion
Calathidius
Calathus
A Genus in the Kingdom Animalia.[1] [more]
Calleida
Callida
Callisthenes
A Genus in the Kingdom Animalia.[2] [more]
Callistoides
Callistomimus
Callistus
Callistus, Callixtus (both Latin) and Kallistos (in Greek) may refer to: [more]
Callytron
Calodromius
Calomera
Calophaena
Calosama
Calosoma
Calybe
Campalita
Canarobius
Carabites
Carabomorphus
Carabus
Carabus is a genus of beetle in family Carabidae. [more]
Cardiaderus
Cardiomera
Carenum
Cassolaia
Castelnaudia
Catadromus
Catascopus
Cedrorum
Celaenephes
Cephalota
Ceroglossus
Cerotalis
Chaetocarabus
Chaetodera
Chalcomenus
Cheporus
Chinajujiroa
Chlaeniellus
Chlaeniomimus
Chlaenius
Chrysocarabus
Chylnus
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
Cillenus
Clarencia
Clivina
Clivinia
Colfax
Colliuris
Colpocaccus
Colpodes
Colpodiscus
Conopterum
Coptocarpus
Coptodera
Coptodere
Coptolabrus
Corsyra
Cosmodela
Cosmodiscus
Crasodactylus
Craspedonotus
Craspedophorus
Cratacanthus
Cratoferonia
Cratogaster
Croatotrechus
Cryptophonus
Ctenosta
Ctenostoma
Cuneipectus
Curtonotus
Cychrus
Cyclotrachelus
Cylindera
Cylindronotum
Cymbionotum
Cymindis
Cypholoba
Dalmataphaenops
Dalyat
Damaster
Daptus
Darlingtonea
Darodilia
Decogmus
Delichoctis
Delinius
Deltomerus
Demetrias
A Genus in the Kingdom Animalia. [more]
Demetrida
Dercylinus
Derobroscus
Deropristus
Desera
Dhanya
Diacheila
Diachromus
Dicaelus
Dicheirotrichus
Dicheirus
Dichrochile
Dicranoglossus
Dicranoncus
Dicranonus
Dicraspeda
Dicrodontus
Dineutes
Dinodes
Diplochaetus
Diplocheila
Diplochelia
Diplous
Disamara
Dischissus
Discoderus
Disenochus
Distichus
Distipsidera
Ditomus
Dixus
Doderotrechus
Dolichoctis
Dolichus
Dolischus
Drimostoma
Dromochorus
Drypta
Duvaliopsis
Duvalius
Dyschiriodes
Dyschirius
Dyschirus
Dystrichothorax
Eccoptomenus
Eccoptoptera
Ega
A Genus in the Kingdom Animalia.[3] [more]
Egadroma
Elaphropus
Elaphrus
Emphanes
Enceladus
Endynomena
Enkelgezum
Entomoantyx
Eobroscus
Eocarterus
Eochlaenius
Epaphiopsis
Epaphius
Epilectus
Episcopellus
Epomis
Eripus
Eucaerus
Eucarterus
Eucolliuris
Eudalia
Eugrapha
Euphorticus
Euplynes
Euproctinus
Europhilus
Euryaptus
Eurycarabus
Euryderus
Eurylychnus
Eurylynchus
Eurynebria
Euryscaphus
Eurysoma
Eurystomis
Euschizomerus
Eusthulmannium
Euthenarus
Eutoma
Eutrechopsis
Eutrechus
Eutrichopus
Evolenes
Formosiella
Fortagonum
Galaicodytes
Galerita
Galeritula
Galiciotyphlotes
Gastrellarius
Gatrellarius
Gehringia
Geobaenus
Geopinus
Geoscaptus
Gigadema
Glabrotrechus
Gnathaphanus
Gnatholymnaeum
Gnathoxys
Goniotropus
Gonogenia
Graniger
Graphipterus
Gynandromorphus
Gynandropus
Habrodera
Hadrocarabus
Halocoryza
Haolochlaenius
Haplaner
Haplochiaenius
Haplochlaenius
Harpalellus
Harpalobrachys
Harpalus
A Genus in the Kingdom Animalia.[4] [more]
Harplaner
Hartonymus
Helluapterus
Helluarchus
Helluodema
Helluomorphoides
Helluonidius
Hemicarabus
Henrotius
Hiletus
Hispalis
Holcoderus
Hololeius
Homethes
Homodela
Hormacrus
Horologion
A Genus in the Kingdom Animalia. [more]
Hydraphaenops
Hydrotrechus
Hygrocarabus
Hyparpalus
Hyperion
Hypharpax
Hypotyphlus
Iberanillus
Iberodytes
Iberotrechus
Idacarabus
Iga
Ildobates
Illaphanus
Infernophilus
Inna
Ishidatrechus
Italaphaenops
Italodytes
Jdiochroma
Jujiroa
Laccocenus
Laccopterum
Laccoscaphus
Lachnocrepis
Lachnoderma
Lachnolebia
Lachnophorus
Lacordairia
Laemostenus
Lamprias
Lasiotrechus
Lauosterus
Lebia
Lebidia
Lecalida
Lecanomerus
Leiradira
Leistus
Leptocarabus
Lessinodytes
Lesticus
Lestignathus
Licinopsis
Licinus
Limnastis
Limodromus
Lindrothius
Lionychus
Lioptera
Lipaster
Lissopogonus
Lithochlaenius
Lonchosternus
Lophoglossus
Lophyra
Lophyridia
Loricera
Lorocera
Lorostema
Lovricia
Loxandrus
Loxodactylus
Loxogenius
Loxoncus
Luperca
Lymnaeum
Lymnastis
Lymnastus
Lymneops
Mantichora
Masoreus
Mauna
Mecodema
Mecodema is a genus of ground beetle in the family Carabidae. [more]
Mecomenus
Mecyclothorax
Mecynognathus
Megacephala
Megalopaussus
Melisodera
Meonis
Mesolestes
Mesothriscus
Metabletus
Metacolpodes
Metadromius
Metallina
Metrius
Metromenus
Micralymma
A Genus in the Kingdom Animalia. [more]
Micratopus
Micrixys
Microcosmodes
Microderes
Microferonia
Microleser
Microlestes
Microlestodes
Microtyphlus
Migadops
Mimotrechus
Minypatrobus
Mioptachys
Miscelus
Miscodera
Molopidius
Molops
Monocentrum
Morimotoidius
Moriodema
Morion
A Genus in the Kingdom Animalia.[5] [more]
Moriosomus
Mormolyce
Mouhotia
Muscelus
Myas
Myriochila
Myriochile
Myrmecodemus
Mysticomenus
Mystropomus
Nabria
Neaphaenops
Nebria
Nelsonites
Nemotarsus
Neocarenum
Neoeudalia
Neohelluo
Neolaphyra
Neonomius
Neoscaphus
Neotrechus
Nesacinopus
Nesorthomus
Netrodera
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
Notaphus
Nothiophilus
Notiobia
Notiophilus
Notonomus
Notoplatynus
Notospeophonus
Nototarsus
Nototrechus
Nurus
Ocyropus
Ocys
Odacantha
Odacanthella
Odacentha
Odontium
Odontocarus
Odontocheila
Odontonyx
Oedesis
Ohomopterus
Oliskhopus
Olisthopus
Omophron
Omphra
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
Ophionea
Ophonus
Opisthius
Oreonebria
Orinicarabus
Orinocarabus
Orionella
Orotrechus
Orthogonius
Orthomus
Orzolina
Oscadytes
Oxycentrus
Oxycheila
Oxycrepis
Oxydrepanus
Oxyglychus
Oxygonia
Oxypselaphus
Ozaena
A Genus in the Kingdom Animalia. [more]
Pachydinodes
Pachyteles
Pamborus
Panagaeus
Pangus
Parabroscus
Paradromius
Paraeutrichopus
Parallelomorphus
Paranchus
Paraphaenops
Paratachys
Parazuphium
Parena
Parophonus
Parroa
Pasimachus
Passalidius
Patrobus
Paussus
Pediomorphus
Pedius
Pelmatellus
Pelophila
Penetretus
Pentacomia
Pentagonica
Percosia
Percosoma
Percus
Pericalus
Pericompsus
Pericompus
Perigona
Perileptus
Peripristus
Peripristus ater is a species of beetle in the family Carabidae, the only species in the genus Peripristus. [more]
Peronomerus
Pheggomisetes
Pheropsophus
Philochthus
Philorhizus
Philoscaphus
Phloeoxena
Phloeozeteus
Phorticosomus
Phrypeus
Physea
Piesmus
Pinacodera
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
Platus
Platycarabus
Platycerozaena
Platyderus
Platylytron
Platynus
Platypatrobus
Platysma
A Genus in the Kingdom Animalia. [more]
Platytarus
Plochinus
Plochionus
Poecilus
Pogonistes
Pogonodaptus
Pogonoglossus
Pogonus
Poliocypas
Polistichus
Polpochila
Polyderis
Polystichus
Porotachys
Prena
Princidium
Pristonychus
Pristosia
Procarabus
Procerus
A Genus in the Kingdom Animalia.[6] [more]
Procrustes
A Genus in the Kingdom Animalia. [more]
Procustes
Prodisenochus
Promecoderus
Promecognathus
Prosopogmus
Pseudabarys
Pseudanophthalmus
Pseudaptinus
Pseudobroscus
Pseudomasoreus
Pseudomorpha
Pseudomyas
Pseudoophonus
Pseudoopterus
Pseudotaphoxenus
Pseudotrechus
Pseudoxycheila
Psydrus
Pterostichus
Quammenis
Rambousekiella
Reicheia
Reicheiodes
Rembus
Rhadine
Rhegmatobius
Rhopalostyla
Rhytisternus
Risophilus
Rupa
Sakagutia
Sardaphaenops
Sarothrocrepis
Sarticus
Scambocarabus
Scaphinotus
Scaraphites
Scarites
Schizogenius
Scopodes
Scotodipnus
Scotoplanetes
Scybalicus
Selenophorus
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
Somoplatus
Somotrichus
Spelaeodytes
Spelaeovulcania
Speleotyphlus
Speluncarius
Speomolops
Speotrechus
Sphaeroderus
Sphaerotachys
Sphallomorpha
Spheroderus
Sphodropsis
Sphodrus
Stenocrepis
Stenolophus
Stenomorphus
Stereocerus
Stichonotus
Stolonis
Stomis
Stricklandia
Stylulus
Styracoderus
Synchus
Syntomus
Synuchidius
Synuchus
Tachys
Tachyta
Tachyura
Taicona
Tanystoma
Tanythrix
Taphoxenus
Tapinopterus
Tasmanitachoides
Tecnophilus
Tefflus
Tetracha
Tetragonoderus
Tetraleucus
Thalassophilus
Thalassotrechus
Thalpius
22503 Thalpius (1997 TB12) is a Jupiter Trojan discovered on October 7, 1997 by M. Tichy and Z. Moravec at Klet. [more]
Therates
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
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
Trechicus
Trechimorphus
Trechoblemus
Trechus
Trepanes
Trephionus
Trichaphaenops
Trichis
Trichocellus
Trichosternus
Trichotichnus
Tricondyla
Trigonognatha
Trigonotoma
Tritrichis
Troglanillus
Troglorites
Trymosternus
Tschitscherinellus
Typhlocharis
Typhlochoromus
Typhlonesiotes
Typhloreicheia
Typhlotrechus
Typsiharpalus
Vachinius
Wolltinerfia
Xenion
Xenotrechus
Xestonotus
Zabrus
Zacotus
Zargus
Zariquieya
Zecillenus
Zeodera
Zuphium
At least 14 species and subspecies belong to the Genus Zuphium.
More info about the Genus Zuphium may be found here.
References
- ^ "Carabidae Taxa". Carabidae of the World. 2011. http://carabidae.pro/carabidae. Retrieved 24 Jun 2011.
- ^ 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.
- ^ John L. Capinera. Encyclopedia of Entomology. p. 1746.
- ^ 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.
- ^ "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.
- ^ 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.
- ^ Charles Darwin (1846). "Letter to Leonard Jenyns, October 17, 1846". http://www.darwinproject.ac.uk/darwinletters/calendar/entry-1009.html.
- ^ Shozo Osawa, Zhi-Hui Su & Yuki Inmura (2004). Molecular Phylogeny and Evolution of Carabid Ground Beetles. Springer. ISBN 4-431-00487-4.
- ^ I. L?bl & ; A. Smetana, ed. (2003?). Catalogue of Palaearctic Coleoptera. Stenstrup, Denmark: Apollo Books.
- ^ "Trees of family Carabidae". Carabidae of the World Database. 2008. http://www.carabidae.ru/carabidae.htm. Retrieved July 24, 2008.
- ^ 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.
- ^ 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.
- ^ "Harpalinae". Fauna Europaea. 2004. http://www.faunaeur.org/full_results.php?id=379457. Retrieved February 16, 2011.
- ^ 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
- http://bugguide.net/index.php?q=search&keys=Calathus&search=Search
- http://bugguide.net/index.php?q=search&keys=Callisthenes&search=Search
- http://bugguide.net/index.php?q=search&keys=Ega&search=Search
- http://bugguide.net/index.php?q=search&keys=Harpalus&search=Search
- http://bugguide.net/index.php?q=search&keys=Morion&search=Search
- http://bugguide.net/index.php?q=search&keys=Procerus&search=Search
- ^ 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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