Apocrita is a
suborder
Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
of
insects in the order
Hymenoptera
Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic.
Females typica ...
. It includes
wasps,
bee
Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...
s, and
ants, and consists of many families. It contains the most advanced hymenopterans and is distinguished from
Symphyta
Sawflies are the insects of the suborder Symphyta within the order Hymenoptera, alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor, which the females use to cut into the plants where they lay ...
by the narrow "waist" (
petiole) formed between the first two segments of the actual
abdomen; the first abdominal segment is fused to the thorax, and is called the
propodeum
The propodeum or propodium is the first abdominal segment in Apocrita Hymenoptera ( wasps, bees and ants). It is fused with the thorax to form the mesosoma. It is a single large sclerite, not subdivided, and bears a pair of spiracles. It is ...
. Therefore, it is general practice, when discussing the body of an apocritan in a technical sense, to refer to the
mesosoma and
metasoma (or
gaster) rather than the "thorax" and "abdomen", respectively. The evolution of a constricted waist was an important adaption for the
parasitoid
In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
lifestyle of the ancestral apocritan, allowing more maneuverability of the female's
ovipositor
The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
. The
ovipositor
The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
either extends freely or is retracted, and may be developed into a stinger for both defense and paralyzing prey. Larvae are legless and blind, and either feed inside a host (plant or animal) or in a nest cell provisioned by their mothers.
Apocrita has historically been split into two groups,
Parasitica and
Aculeata. Aculeata is a
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term ...
whose name is in standard use. "Parasitica" is not a clade, as it is
paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
: the clade would contain the Aculeata. "Parasitica" is therefore a rankless grouping in many present classifications, if it appears at all.
Parasitica comprises the majority of hymenopteran insects, its members living as
parasitoids
In evolutionary ecology, a parasitoid is an organism that lives in close association with its host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionary strategies within parasitis ...
.
Most species are small, with the ovipositor adapted for piercing. In some hosts, the parasitoids induce metamorphosis prematurely, and in others it is prolonged. There are even species that are hyperparasites, or parasitoids on other parasitoids. The Parasitica lay their egg
An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
s inside or on another insect (egg, larva or pupa) and their larvae grow and develop within or on that host. The host is nearly always killed. Many parasitic hymenopterans are used as biological control
Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically als ...
agents to control pests, such as caterpillar
Caterpillars ( ) are the larva, larval stage of members of the order Lepidoptera (the insect order comprising butterfly, butterflies and moths).
As with most common names, the application of the word is arbitrary, since the larvae of sawfly ...
s, true bugs and hoppers, flies
Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwing ...
, and weevil
Weevils are beetles belonging to the superfamily Curculionoidea, known for their elongated snouts. They are usually small, less than in length, and herbivorous. Approximately 97,000 species of weevils are known. They belong to several families, ...
s.
Aculeata is a monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...
group that includes those species in which the female's ovipositor
The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
is modified into a stinger
A stinger (or sting) is a sharp organ found in various animals (typically insects and other arthropods) capable of injecting venom, usually by piercing the epidermis of another animal.
An insect sting is complicated by its introduction of ve ...
to inject venom
Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a ...
. Groups within Aculeata include the familiar ants, bees, and various types of parasitic and predatory wasps; it also includes all of the social
Social organisms, including human(s), live collectively in interacting populations. This interaction is considered social whether they are aware of it or not, and whether the exchange is voluntary or not.
Etymology
The word "social" derives from ...
hymenopterans.
Among the nonparasitic and nonsocial Aculeata, larvae are fed with captured prey (typically alive and paralyzed) or may be fed pollen and nectar. The social
Social organisms, including human(s), live collectively in interacting populations. This interaction is considered social whether they are aware of it or not, and whether the exchange is voluntary or not.
Etymology
The word "social" derives from ...
Aculeata feed their young prey ( paper wasps and hornet
Hornets (insects in the genus ''Vespa'') are the largest of the eusocial wasps, and are similar in appearance to their close relatives yellowjackets. Some species can reach up to in length. They are distinguished from other vespine wasps by the ...
s), or pollen
Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...
and nectar (bees), or perhaps seeds, fungi
A fungus (plural, : fungi or funguses) is any member of the group of Eukaryote, eukaryotic organisms that includes microorganisms such as yeasts and Mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified ...
, or nonviable eggs (ants).
Extant families and superfamilies
The Apocrita contains a large number of families. Some traditional taxa such as the Parasitica (containing many families of parasitoid wasp
Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps ( Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later cau ...
s) have been found on molecular analysis to be paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
. Parasitoidism evolved once, and it is found today across most Apocritan families, though it has been secondarily lost several times. The phylogenetic tree gives a condensed overview of the phylogeny, illustrated with major groups. The sawflies are paraphyletic as the Apocrita evolved inside that group. The tree is not fully resolved.
*Suborder Apocrita
**Infraorder Aculeata
***Superfamily Apoidea
The superfamily Apoidea is a major group within the Hymenoptera, which includes two traditionally recognized lineages, the "sphecoid" wasps, and the bees. Molecular phylogeny demonstrates that the bees arose from within the traditional "Crabronida ...
(bee
Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...
s and sphecoid wasps)
****Family Ampulicidae (cockroach wasp
The Ampulicidae, or cockroach wasps, are a small (about 170 species), primarily tropical family of sphecoid wasps, all of which use various cockroaches as prey for their larvae. They tend to have elongated jaws, pronounced neck-like constrictions ...
s)
****Family Andrenidae
The Andrenidae (commonly known as mining bees) are a large, nearly cosmopolitan family of solitary, ground-nesting bees. Most of the family's diversity is located in temperate or arid areas (warm temperate xeric). It includes some enormous gene ...
(mining bees)
****Family Apidae
Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen bees, including bumblebees and honey bees, but also includes stingless bees (also used ...
(carpenter bee
Carpenter bees are species in the genus ''Xylocopa'' of the subfamily Xylocopinae. The genus includes some 500 bees in 31 subgenera. The common name "carpenter bee" derives from their nesting behavior; nearly all species burrow into hard plant m ...
s, digger bees, cuckoo bee
The term cuckoo bee is used for a variety of different bee lineages which have evolved the kleptoparasitic behaviour of laying their eggs in the nests of other bees, reminiscent of the behavior of cuckoo birds. The name is perhaps best applied to ...
s, bumble bee
A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related genera ...
s, orchid bees, stingless bee
Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina according to other authors). They belong in the family ...
s, and honeybee
A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cos ...
s)
****Family Colletidae (yellow-faced bee
The Colletidae are a family of bees, and are often referred to collectively as plasterer bees or polyester bees, due to the method of smoothing the walls of their nest cells with secretions applied with their mouthparts; these secretions dry int ...
s and plasterer bees)
****Family Crabronidae
The Crabronidae are a large paraphyletic group (nominally a family) of wasps, including nearly all of the species formerly comprising the now-defunct superfamily Sphecoidea. It collectively includes well over 200 genera, containing well over ...
(sand wasps, bee wolves, etc.)
****Family Halictidae
Halictidae is the second-largest family of bees (clade Anthophila) with nearly 4,500 species. Halictid species are an extremely diverse group that can vary greatly in appearance. These bees occur all over the world and are found on every contine ...
("sweat bees")
****Family Heterogynaidae
****Family Megachilidae
Megachilidae is a cosmopolitan family of mostly solitary bees. Both that their pollen-carrying structure (called a '' scopa'') is restricted to the ventral surface of the abdomen (rather than mostly or exclusively on the hind legs as in other ...
( leaf-cutting bees)
****Family Melittidae
****Family Stenotritidae
****Family Sphecidae ( digger wasps)
***Superfamily Chrysidoidea
****Family Bethylidae
The Bethylidae are a family of aculeate wasps in the superfamily Chrysidoidea. As a family, their biology ranges between parasitoid wasps and hunting wasps.
Overview
Like most of the Chrysidoidea, the Bethylidae are stinging Hymenoptera and ...
****Family Chrysididae (cuckoo wasp
Commonly known as cuckoo wasps or emerald wasps, the hymenopteran family Chrysididae is a very large cosmopolitan group (over 3000 described species) of parasitoid or kleptoparasitic wasps, often highly sculptured, with brilliant metallic col ...
s)
****Family Dryinidae
****Family Embolemidae
****Family Plumariidae
****Family Sclerogibbidae
****Family Scolebythidae
***Superfamily Formicoidea
****Family Formicidae ( ants)
***Superfamily Pompiloidea
****Family Mutillidae ( velvet ants)
****Family Myrmosidae
****Family Pompilidae
Wasps in the family Pompilidae are commonly called spider wasps, spider-hunting wasps, or pompilid wasps. The family is cosmopolitan, with some 5,000 species in six subfamilies. Nearly all species are solitary (with the exception of some group-ne ...
(spider wasp
Wasps in the family Pompilidae are commonly called spider wasps, spider-hunting wasps, or pompilid wasps. The family is cosmopolitan, with some 5,000 species in six subfamilies. Nearly all species are solitary (with the exception of some group- ...
s)
****Family Sapygidae
The Sapygidae are a family of solitary kleptoparasitic aculeate wasps. They are generally black wasps, similar in appearance to some Tiphiidae or Thynnidae, with white or yellow markings developed to various degrees.
The female oviposits her ...
***Superfamily Scolioidea
****Family Scoliidae
***Superfamily Tiphioidea
****Family Bradynobaenidae
****Family Sierolomorphidae
****Family Tiphiidae
***Superfamily Thynoidea
****Family Chyphotidae
****Family Thynnidae
***Superfamily Vespoidea
****Family Rhopalosomatidae
****Family Vespidae
The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', '' Vespa orientalis'', and '' Vespula germanica'') and many solitary wasps. Ea ...
( paper wasps, potter wasps, hornet
Hornets (insects in the genus ''Vespa'') are the largest of the eusocial wasps, and are similar in appearance to their close relatives yellowjackets. Some species can reach up to in length. They are distinguished from other vespine wasps by the ...
s, pollen wasp
Pollen wasps, the Masarinae, are unusual wasps that are typically treated as a subfamily of Vespidae, but have in the past sometimes been recognized as a separate family, "Masaridae", which also included the subfamilies Euparagiinae and Gayellinae ...
s, yellowjacket
Yellowjacket or yellowjacket is the common name in North America for predatory social wasps of the genus, genera ''Vespula'' and ''Dolichovespula''. Members of these genera are known simply as "wasps" in other English-speaking countries. Most of ...
s)
**Infraorder Parasitica
***Superfamily Ceraphronoidea
****Family Ceraphronidae
****Family Megaspilidae
***Superfamily Chalcidoidea
Chalcid wasps (, , for their metallic colour) are insects within the superfamily Chalcidoidea, part of the order Hymenoptera. The superfamily contains some 22,500 known species, and an estimated total diversity of more than 500,000 species, m ...
****Family Agaonidae
The family Agaonidae is a group of pollinating and nonpollinating fig wasps. They spend their larval stage inside the fruits of figs. The pollinating wasps ( Agaoninae, Kradibiinae, and Tetrapusiinae) are the mutualistic partners of the fig tr ...
(fig wasp
Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside figs. Most are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, while ...
s)
****Family Aphelinidae
The Aphelinidae are a moderate-sized family of tiny parasitic wasps, with about 1100 described species in some 28 genera. These minute insects are challenging to study, as they deteriorate rapidly after death unless extreme care is taken (e.g., p ...
****Family Chalcididae (chalcid wasp
Chalcid wasps (, , for their metallic colour) are insects within the superfamily Chalcidoidea, part of the order Hymenoptera. The superfamily contains some 22,500 known species, and an estimated total diversity of more than 500,000 species, me ...
s)
****Family Encyrtidae
****Family Eucharitidae
****Family Eulophidae
The Eulophidae are a large family of hymenopteran insects, with over 4,300 described species in some 300 genera. The family includes the genus '' Elasmus'', which used to be treated as a separate family, "Elasmidae", and is now treated as a su ...
****Family Eupelmidae
****Family Eurytomidae
The Eurytomidae are a family within the superfamily Chalcidoidea.
Unlike most chalcidoids, the larvae of many are phytophagous (feeding in stems, seeds, or galls), while others are more typical parasitoids, though even then the hosts are usually ...
(seed chalcid
The Eurytomidae are a family within the superfamily Chalcidoidea.
Unlike most chalcidoids, the larvae of many are phytophagous (feeding in stems, seeds, or galls), while others are more typical parasitoids, though even then the hosts are usual ...
s)
****Family Leucospidae
The Leucospidae (sometimes incorrectly spelled Leucospididae) are a specialized group of wasps within the superfamily Chalcidoidea, that are ectoparasitoids of aculeate wasps or bees. They are typically mimics of bees or stinging wasps, often ...
****Family Mymaridae ( fairyflies) – the smallest of all insects
****Family Ormyridae
****Family Perilampidae
****Family Pteromalidae
The Pteromalidae are a very large family of mostly parasitoid wasps, with some 3,450 described species in about 640 genera (the number was greater, but many species and genera have been reduced by synonymy in recent years). The subfamily-level di ...
****Family Rotoitidae
The Rotoitidae are a very small family of rare, relictual parasitic wasps in the superfamily Chalcidoidea, known primarily from fossils (14 extinct species in two genera, '' Baeomorpha'' and '' Taimyromorpha'').Huber JT, Shih C, Dong R (2019) A n ...
****Family Signiphoridae
Signiphoridae (historically also known as Thysanidae) is a small family of parasitic wasps in the superfamily Chalcidoidea. The roughly 80 species are placed in four genera.
Diagnosis
The Signiphoridae range in size from . They are usually blac ...
****Family Tanaostigmatidae
The Tanaostigmatidae are a small family of parasitic wasps in the superfamily Chalcidoidea. They are almost exclusively phytophagy, phytophagous insects, forming galls in plant stems, leaves, or seeds. The some 90 species in 9 genera are primaril ...
****Family Tetracampidae
The Tetracampidae are a small family of parasitic wasps in the superfamily Chalcidoidea. They are parasitoids of phytophagous insects, primarily flies. The 44 species in 15 genera are almost entirely absent from the New World
The term ''N ...
****Family Torymidae
****Family Trichogrammatidae
The Trichogrammatidae are a family of tiny wasps in the Chalcidoidea
Chalcid wasps (, , for their metallic colour) are insects within the superfamily Chalcidoidea, part of the order Hymenoptera. The superfamily contains some 22,500 known ...
***Superfamily Cynipoidea
****Family Austrocynipidae
****Family Cynipidae
Gall wasps, also incorrectly called gallflies, are hymenopterans of the family Cynipidae in the wasp superfamily Cynipoidea. Their common name comes from the galls they induce on plants for larval development. About 1,300 species of this gener ...
(gall wasp
Gall wasps, also incorrectly called gallflies, are hymenopterans of the family Cynipidae in the wasp superfamily Cynipoidea. Their common name comes from the galls they induce on plants for larval development. About 1,300 species of this generall ...
s)
****Family Figitidae
****Family Ibaliidae
****Family Liopteridae
***Superfamily Diaprioidea
****Family Austroniidae
****Family Diapriidae
The Diapriidae are a family of parasitoid wasps. These tiny insects have an average length of 2–4 mm and never exceed 8 mm. They typically attack larvae and pupae of a wide range of insects, especially flies. The about 2,300 described ...
****Family Maamingidae
****Family Monomachidae
***Superfamily Evanioidea
The Evanioidea are a small hymenopteran superfamily that includes three families, two of which (Aulacidae and Gasteruptiidae) are much more closely related to one another than they are to the remaining family, Evaniidae. The rich fossil record, h ...
****Family Aulacidae
****Family Evaniidae (ensign wasp
Evaniidae is a family of parasitoid wasps also known as ensign wasps, nightshade wasps, hatchet wasps, or cockroach egg parasitoid wasps. They number around 20 extant genera containing over 400 described species, and are found all over the world e ...
s)
****Family Gasteruptiidae
***Superfamily Ichneumonoidea
The superfamily Ichneumonoidea contains one extinct and three extant families, including the two largest families within Hymenoptera: Ichneumonidae and Braconidae. The group is thought to contain as many as 100,000 species, many of which have not ...
****Family Braconidae
The Braconidae are a family of parasitoid wasps. After the closely related Ichneumonidae, braconids make up the second-largest family in the order Hymenoptera, with about 17,000 recognized species and many thousands more undescribed. One analy ...
****Family Ichneumonidae
The Ichneumonidae, also known as the ichneumon wasps, Darwin wasps, or ichneumonids, are a family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 25,000 species cu ...
(ichneumon wasp
The Ichneumonidae, also known as the ichneumon wasps, Darwin wasps, or ichneumonids, are a family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 25,000 species cu ...
s)
***Superfamily Megalyroidea
****Family Megalyridae
Megalyroidea is a small hymenopteran superfamily of wasps that includes a single family, Megalyridae, with eight extant genera (plus around a dozen extinct ones) and 49 described species. Modern megalyrids are found primarily in the southern hem ...
***Superfamily Mymarommatoidea – sometimes called Serphitoidea
Proctotrupomorpha is a major subgrouping of the Apocrita within the Hymenoptera, containing mainly parasitic wasps. It contains the major groupings of Chalcidoidea, Diaprioidea, Proctotrupoidea, Cynipoidea and Platygastroidea, as well as the sma ...
****Family Mymarommatidae
***Superfamily Platygastroidea
The Hymenopteran superfamily of parasitoid wasps, Platygastroidea, has often been treated as a lineage within the superfamily Proctotrupoidea, but most classifications since 1977 have recognized it as an independent group within the Proctotrupomor ...
****Family Platygastridae
The hymenopteran family Platygastridae (sometimes incorrectly spelled Platygasteridae) is a large group (over 4000 species) of exclusively parasitoid wasps, mostly very small (1–2 mm), black, and shining, with geniculate (elbowed) anten ...
****Family Scelionidae
The hymenopteran family Scelionidae is a very large cosmopolitan group (over 3000 described species in some 176 genera) of exclusively parasitoid wasps, mostly small (0.5–10 mm), often black, often highly sculptured, with (typically) elbowe ...
***Superfamily Proctotrupoidea
Proctotrupoidea is a hymenopteran superfamily containing seven extant families, though others have been recognized in the past, most of these having been removed to a recently erected superfamily Diaprioidea
Diaprioidea is a hymenopteran supe ...
****Family Heloridae
****Family Pelecinidae
****Family Peradeniidae
****Family Proctorenyxidae
****Family Proctotrupidae
Proctotrupidae is a family of wasps in the superfamily Proctotrupoidea of the order Hymenoptera. There are about 400 species in more than 30 genera in Proctotrupidae, found throughout most of the world.
Proctotrupidae are small parasitoid wasps, ...
****Family Roproniidae
Roproniidae is a family of wasps in the order Hymenoptera, of which only two genera are still extant, the others being fossils.
Extant genera
* '' Ropronia'' Provancher, 1887
* '' Xiphyropronia'' He & Chen, 1991
Fossil genera
* † '' Beipiaosi ...
****Family Vanhorniidae
***Superfamily Stephanoidea
Stephanoidea is a superfamily of parasitic wasps within the Apocrita, it includes only one living family, Stephanidae (350 living species mid Cretaceous-recent), as well as the extinct families Ephialtitidae (89 species, Early Jurassic-mid Cretac ...
****Family Stephanidae
The Stephanidae, sometimes called crown wasps, are a family of parasitoid wasps. They are the only living members of the superfamily Stephanoidea. Stephanidae has at least 345 living species in 11 genera. The family is considered cosmopolitan in ...
***Superfamily Trigonaloidea
Trigonalidae (the spelling Trigonalyidae is incorrect under Article 29.5 of the ICZN,[ICZN Code](_blank)
29.5. Maintena ...
****Family † Maimetshidae
****Family Trigonalidae
References
External links
Suborder Apocrita – Ants, Bees and Wasps – BugGuide.Net
— images and other information
Tree of Life
Balades Entomologiques
— "entomological walks" with images
{{Taxonbar, from=Q22864
Insect suborders
Taxa named by Carl Eduard Adolph Gerstaecker