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The Pyralidae, commonly called pyralid moths, snout moths or grass moths, are a
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
in the ditrysian
superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
Pyraloidea The Pyraloidea (pyraloid moths or snout moths) are a moth superfamily containing about 16,000 described species worldwide, and probably at least as many more remain to be described. They are generally fairly small moths, and as such, they have b ...
. In many (particularly older) classifications, the grass moths (Crambidae) are included in the Pyralidae as a
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
, making the combined group one of the largest families in the Lepidoptera. The latest review by
Eugene G. Munroe Eugene Gordon Munroe (8 September 1919 – 31 May 2008) was a Canadian entomologist who discovered numerous species of insects. He worked for the Insect Systematics and Biological Control Unit, Entomology Division in Ottawa, Ontario, Canada. Munro ...
and
Maria Alma Solis Maria Alma Solis is a entomologist at the Systematic Entomology Laboratory (SEL) of the Agricultural Research Service of the United States Department of Agriculture, U.S. Department of Agriculture. Early life Maria Alma Solis was born on Feb ...
retain the Crambidae as a full family of
Pyraloidea The Pyraloidea (pyraloid moths or snout moths) are a moth superfamily containing about 16,000 described species worldwide, and probably at least as many more remain to be described. They are generally fairly small moths, and as such, they have b ...
. The
wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingspan of ...
s for small and medium-sized species are usually between with variable morphological features. It is a diverse group, with more than 6,000 species described worldwide, and more than 600 species in America north of Mexico, comprising the third largest moth family in North America. At least 42 species have been recorded from North Dakota in the subfamilies of Pyralidae.


Relationship with humans

Most of these small moths are inconspicuous. Many are economically important pests, including waxworms, which are the
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e of the greater ('' Galleria mellonella'') and lesser (''
Achroia grisella The lesser wax moth (''Achroia grisella'') is a small moth of the snout moth family (Pyralidae) that belongs to the subfamily Galleriinae. The species was first described by Johan Christian Fabricius in 1794. Adults are about 0.5 inches (13  ...
'') wax moths (
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
Galleriinae The Galleriinae are a subfamily of snout moths (family Pyralidae) and occur essentially worldwide, in some cases aided by involuntary introduction by humans. This subfamily includes the wax moths, whose caterpillars (waxworms) are bred on a comme ...
). They are natively
pest Pest or The Pest may refer to: Science and medicine * Pest (organism), an animal or plant deemed to be detrimental to humans or human concerns ** Weed, a plant considered undesirable * Infectious disease, an illness resulting from an infection ** ...
s of beehives, but are bred indoors in enormous numbers as live food for small
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
and
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
pets and similar animals. They are also used as fishing bait for
trout Trout are species of freshwater fish belonging to the genera '' Oncorhynchus'', ''Salmo'' and ''Salvelinus'', all of the subfamily Salmoninae of the family Salmonidae. The word ''trout'' is also used as part of the name of some non-salmoni ...
fishing. Other notable snout moth pests relevant for their larval hosts include: * Alligatorweed stem borer (''
Arcola malloi ''Arcola malloi'' (formerly ''Vogtia malloi'') is a species of snout moth known as the alligator weed stem borer. It is used as an agent of biological pest control against the noxious aquatic plant known as alligator weed (''Alternanthera philox ...
'': Phycitinae) – biological control of alligator weed (''
Alternanthera philoxeroides ''Alternanthera philoxeroides'', commonly referred to as alligator weed, is a native species to the temperate regions of South America, which includes Argentina, Brazil, Paraguay and Uruguay. Argentina alone hosts around 27 species that fall with ...
''). * Almond moth (''
Cadra cautella The almond moth or tropical warehouse moth (''Cadra cautella'') is a small, stored-product pest. Almond moths infest flour, bran, oats, and other grains, as well as dried fruits. It belongs to the family of snout moths (Pyralidae), and more spec ...
'': Phycitinae) – pest of stored
cereal A cereal is any Poaceae, grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, Cereal germ, germ, and bran. Cereal Grain, grain crops are grown in greater quantit ...
s and dry fruit; now introduced almost worldwide. * Cacao moth, tobacco moth, warehouse moth (''
Ephestia elutella ''Ephestia elutella'', the cacao moth, tobacco moth or warehouse moth, is a small moth of the family Pyralidae. It is probably native to Europe, but has been transported widely, even to Australia. A subspecies is ''E. e. pterogrisella''. The w ...
'': Phycitinae) – pest of stored dry vegetable products; Europe, introduced to some other regions (e.g. Australia). * Dried fruit moth (''
Cadra calidella ''Cadra calidella'', the dried fruit or date moth, is a species of snout moth in the genus ''Cadra'' and commonly mistaken for the species ''Cadra figulilella.'' It thrives in warmer conditions and is found primarily in Mediterranean countries, a ...
'': Phycitinae) * ''
Etiella behrii ''Etiella behrii'' is a species of moth of the family Pyralidae. It is found in Hong Kong, Indonesia, Malaysia and most of Australia. The wingspan is about 10 mm. The larvae are considered an agricultural pest on peanut (''Arachis hypogaea ...
'' (Phycitinae) – pest of stored
legume A legume () is a plant in the family Fabaceae (or Leguminosae), or the fruit or seed of such a plant. When used as a dry grain, the seed is also called a pulse. Legumes are grown agriculturally, primarily for human consumption, for livestock f ...
s; Southeast Asia and Australia * " Flour moths" – pests of stored grains, spices, flour, and similar dry vegetable products; now introduced almost worldwide. ** Indian mealmoth (''
Plodia interpunctella The Indianmeal moth (''Plodia interpunctella''), also spelled Indian meal moth and Indian-meal moth, is a pyraloid moth of the family Pyralidae. Alternative common names are weevil moth, pantry moth, flour moth or grain moth. The almond moth ...
'': Phycitinae) ** Mediterranean flour moth, Indian flour moth ('' Ephestia kuehniella'': Phycitinae) * Grease moth ('' Aglossa pinguinalis'':
Pyralinae The Pyralinae are the typical subfamily of snout moths (family Pyralidae) and occur essentially worldwide, in some cases aided by involuntary introduction by humans. They are rather rare in the Americas however, and their diversity in the Austr ...
) – pest of suet and other oily food. * Lesser cornstalk borer (''
Elasmopalpus lignosellus ''Elasmopalpus lignosellus'', the lesser cornstalk borer, was described by Philipp Christoph Zeller in 1848. It is found from the southern United States to Mexico, Central America and South America (Colombia, Venezuela, Brazil, Peru, Argentina and ...
'': Phycitinae) – stalk pest of
corn Maize ( ; ''Zea mays'' subsp. ''mays'', from es, maíz after tnq, mahiz), also known as corn (North American and Australian English), is a cereal grain first domesticated by indigenous peoples in southern Mexico about 10,000 years ago. Th ...
(''Zea mays''); tropical and subtropical Americas, introduced to the Hawaiian Islands. * Locust bean moth ('' Ectomyelois ceratoniae'': Phycitinae) * Mahogany webworm ('' Macalla thyrsisalis'':
Epipaschiinae The Epipaschiinae are a subfamily of snout moths (family Pyralidae). More than 720 species are known today, which are found mainly in the tropics and subtropics. Some occur in temperate regions, but the subfamily is apparently completely absent f ...
) – defoliator pest of mahogany trees (''Swietenia''); Neotropics. * Meal moth (''
Pyralis farinalis ''Pyralis'' is a genus of snout moths. It was described by Carl Linnaeus in 1758. Species * ''Pyralis caustica'' (Meyrick, 1884) * ''Pyralis costinotalis'' Hampson, 1917 * ''Pyralis electalis'' Hulst, 1886 * ''Pyralis farinalis'' (Linnaeus, 175 ...
'': Pyralinae) – pest of stored grain, flour and other cereals; now introduced almost worldwide. * Pear fruit borer ('' Pempelia heringii'': Phycitinae) – pest of apple and pear fruits; East Asia, introduced to the Hawaiian Islands. * Pine webworm (''
Pococera robustella ''Pococera robustella'', the pine webworm moth, is a species of moth of the family Pyralidae. It is found in southern Canada and the eastern United States from Minnesota to New England and south to Florida. The wingspan is 22–25 mm. Adults ...
'': Epipaschiinae) – defoliator pest of
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
s (''Pinus''); North America east of Great Lakes region. * Raisin moth (''
Cadra figulilella ''Cadra figulilella'', the raisin moth, is a moth of the family Pyralidae. The raisin moth is known most commonly as a pest that feeds on dried fruits, such as the raisin and date. It covers a range that includes much of the world, primarily situ ...
'': Phycitinae) – pests of stored dry fruit; now introduced almost worldwide. * Rice moth ('' Corcyra cephalonica'': Galleriinae) – pest of stored grain, flour and other cereals. * South American cactus moth (''
Cactoblastis cactorum ''Cactoblastis cactorum'', the cactus moth, South American cactus moth or nopal moth, is native to Argentina, Paraguay, Uruguay and southern Brazil. It is one of five species in the genus '' Cactoblastis'' that inhabit South America, where many pa ...
'': Phycitinae) – biological control of prickly pears (''Opuntia''). * Southern pine coneworm, "pitch moth" ('' Dioryctria amatella'': Phycitinae) – cone and shoot pest of pines (''Pinus''); southern North America. * Stored nut moth (''
Paralipsa gularis ''Paralipsa gularis'', the stored nut moth, is a moth of the family Pyralidae. It is found in Southeast Asia and is an introduced species in Western Europe. The wingspan is 21–32 mm. The caterpillars feed on stored nuts and seeds like wa ...
'': Galleriinae) – pest of stored nuts and
drupe In botany, a drupe (or stone fruit) is an indehiscent fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pit'', ''stone'', or '' pyrena'') of hardened endocarp with a seed (''kernel'') ...
s; Southeast Asia, introduced to Western Europe. * Sunflower moth (''
Homoeosoma nebulella ''Homoeosoma nebulella'', the Eurasian sunflower moth, is a moth of the family Pyralidae. It is found in Europe, Russia, Anatolia, the Middle East and West Africa. The wingspan is 20–27 mm. The larvae feed on Asteraceae such as ''Cirs ...
'': Phycitinae) – pest of
sunflower seed The sunflower seed is the seed of the sunflower ('' Helianthus annuus''). There are three types of commonly used sunflower seeds: linoleic (most common), high oleic, and sunflower oil seeds. Each variety has its own unique levels of monounsat ...
s; Europe and surrounding regions. The European corn borer (''
Ostrinia nubilalis The European corn borer (''Ostrinia nubilalis''), also known as the European corn worm or European high-flyer, is a moth of the family Crambidae which includes other grass moths. It is a pest of grain, particularly maize (''Zea mays''). The in ...
'') and southern cornstalk borer (''
Diatraea crambidoides ''Diatraea crambidoides'', the southern cornstalk borer moth, is a species of moth of the family Crambidae described by Augustus Radcliffe Grote in 1880. It is found in North America, from Alabama and northern Florida to Ohio and Maryland. Its win ...
''), formerly considered snout moths, are placed in the Crambidae which, as noted above, are usually regarded as a separate family today.


Systematics

Five subfamilies are generally recognized in the Pyralidae today. The Acentropinae (= Nymphulinae), occasionally still placed here, do indeed seem to belong in the Crambidae. The snout moth subfamilies are, listed in the presumed
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
sequence from the most primitive to the most advanced: * Chrysauginae (including Bradypodicolinae, Semniidae) – about 400 species occurring predominantly in the Neotropical region. Larvae typically feed on plants, but some have more unusual feeding habits. The latter include for example some myrmecophilous species, as well as a number of
sloth moth A sloth moth is a coprophagous moth which has evolved to exclusively inhabit the fur of sloths and to use sloth dung as a substrate for the early stages of reproduction. Sloth moths include '' Bradypodicola hahneli'', ''Cryptoses choloepi'', ''Cry ...
s which are dependent on
sloth Sloths are a group of Neotropical xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their li ...
s for their entire life cycle. Most Chrysauginae larvae have a sclerotised ring around seta SD1 of the
metathorax The metathorax is the posterior of the three segments in the thorax of an insect, and bears the third pair of legs. Its principal sclerites ( exoskeletal plates) are the metanotum (dorsal), the metasternum (ventral), and the metapleuron (lateral) ...
. *
Galleriinae The Galleriinae are a subfamily of snout moths (family Pyralidae) and occur essentially worldwide, in some cases aided by involuntary introduction by humans. This subfamily includes the wax moths, whose caterpillars (waxworms) are bred on a comme ...
(including Macrothecinae) – about 300 species worldwide. The males of galleriine moths have a gnathos almost or completely reduced, the pupae have a prominent dorsal median ridge on the thorax and abdomen, and most larvae have a sclerotised ring around seta SD1 of the first abdominal segment. *
Pyralinae The Pyralinae are the typical subfamily of snout moths (family Pyralidae) and occur essentially worldwide, in some cases aided by involuntary introduction by humans. They are rather rare in the Americas however, and their diversity in the Austr ...
(including Endotrichinae, Hypotiinae) – rather diverse in the
Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by the ...
; a lesser number of the roughly 900 species occurs elsewhere. The females of almost all Pyralinae except ''
Cardamyla ''Cardamyla'' is a genus of snout moths. It was described by Francis Walker in 1859 and is known from Australia. Species * ''Cardamyla carinentalis ''Cardamyla carinentalis'' is a species of snout moth in the genus '' Cardamyla''. It was desc ...
'' and ''
Embryoglossa ''Embryoglossa'' is a genus of snout moths. It was described by Warren in 1896, and is known from Nigeria, Madagascar, and India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest cou ...
'' are recognizable by the very short ductus bursae of their genitals. *
Epipaschiinae The Epipaschiinae are a subfamily of snout moths (family Pyralidae). More than 720 species are known today, which are found mainly in the tropics and subtropics. Some occur in temperate regions, but the subfamily is apparently completely absent f ...
(including Pococerinae) – over 550 described species in the tropical and temperate regions (except Europe). Larvae are leaf rollers, leaf tiers, or leaf miners. Some species are minor pests of a few commercial crops. Epipaschiinae are generally hard to recognize, except in the case of adult males which have a few characteristic traits, such as the upturned and pointed third segment of the labial palps and usually a scaly projection from the antenna base. The larvae lack any stereotyped seta sclerotisations. * Phycitinae (including Anerastiinae, Peoriinae) – probably the most difficult group of Pyraloidea in terms of identification and classification. They comprise more than 600 genera and about 4000 species found all over the world. The characteristic trait of the caterpillars is a
sclerotised Sclerotin is a component of the cuticle of various Arthropoda, most familiarly insects. It is formed by cross-linking members of particular classes of protein molecules, a biochemical process called sclerotization, a form of tanning in which qui ...
area encircling the base of seta SD1 on the mesothorax, while the adult females have – like the males of Pyralidae in general do – a frenulum consisting of a single bristle which in turn is composed of multiple acanthae.


Problematic genera

In addition to those assigned to the tribes above, several genera of (presumed) Pyralidae are not firmly placed in this arrangement, but are '' incertae sedis''. Some may be very basal lineages which stand outside the main snout moth radiations, but given the changing circumscription of the Pyralidae, some are likely to be placed outside this group in its modern meaning, either in the Crambidae or in other lineages of basal
Obtectomera The Obtectomera is a clade of macro-moths and butterflies, comprising over 100,000 species in at least 12 superfamilies. Taxonomy The Obtectomera includes the following 12 superfamilies: * Whalleyanoidea Minet, 1991 * Thyridoidea Herrich-Sc ...
. Some may even belong to more ancient moth lineages, such as the Alucitoidea or Pterophoroidea. Finally, some of these (usually little-studied) genera possibly are
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...
s of genera described earlier. These genera are in the unranked category of the family Pyralidae. The genera in question are: * '' Apocabimoides'' Neunzig & Goodson, 1992 * ''
Ardjuna ''Ardjuna'' is a genus of snout moths. It contains the species ''Ardjuna kresna''. It is found on northern Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territor ...
'' Roesler & Küppers, 1979 * '' Cabimoides'' Neunzig & Goodson, 1992 * '' Cryptophycita'' Roesler & Küppers, 1979 * '' Cryptozophera'' Roesler & Küppers, 1979 * '' Delcina'' Clarke, 1986 (Phycitinae?) * '' Eupassadena'' Neunzig & Goodson, 1992 * '' Gomezmenoria'' Agenjo, 1966 * '' Gunungodes'' Roesler & Küppers, 1981 * '' Indocabnia'' Roesler & Küppers, 1981 * '' Inverina'' Neunzig & Goodson, 1992 * '' Kasyapa'' Roesler & Küppers, 1981 * '' Kaurava'' Roesler & Küppers, 1981 * ''
Kumbhakarna Kumbhakarna (Sanskrit: कुम्भकर्ण, lit. ''pot-eared'') is a powerful rakshasa and younger brother of Ravana from the Hindu epic ''Ramayana''. Despite his gigantic size and appetite, he is described as a virtuous character an ...
'' Roesler & Küppers, 1981 * '' Merangiria'' Roesler & Küppers, 1979 * '' Ohigginsia'' Neunzig & Goodson, 1992 * '' Pseudopassadena'' Neunzig & Goodson, 1992 * '' Psorozophera'' Roesler & Küppers, 1979 These genera have been placed in the Pyralidae when these were still circumscribed '' sensu lato'' and are sometimes still treated thus, but actually they seem to belong in the Crambidae (see also ''
Micronix ''Micronix'' is a genus of moths of the family Crambidae. It contains only one species, ''Micronix nivalis'', which is found in Venezuela. References

Crambinae Taxa named by Hans Georg Amsel Crambidae genera {{Crambinae-stub ...
'' and ''
Tanaobela ''Tanaobela'' is a genus of moths in the superfamily Pyraloidea containing only one species, ''Tanaobela chrysochlora'', which is found in Australia, where it has been recorded from Queensland. Its affiliations are disputed. The wingspan Th ...
''): * ''
Alphacrambus ''Alphacrambus'' is a grass moth genus (family Crambidae) of subfamily Crambinae, tribe Crambini. Some authors have placed it in the snout moth The Pyralidae, commonly called pyralid moths, snout moths or grass moths, are a family of Lepidopte ...
'' Bassi, 1995 * ''
Peniculimius ''Peniculimius'' is a grass moth genus (family Crambidae) of subfamily Crambinae; its relationships within the latter group remain obscure. Some authors have placed it in the snout moth The Pyralidae, commonly called pyralid moths, snout moth ...
'' Schoute, 1994 * ''
Steneromene ''Steneromene'' is a grass moth genus (family Crambidae) of subfamily Crambinae, tribe Diptychophorini. Some authors have placed it in the snout moth The Pyralidae, commonly called pyralid moths, snout moths or grass moths, are a family of L ...
'' Gaskin, 1986 * ''
Thopeutis ''Thopeutis'' is a grass moth genus (family Crambidae) of subfamily Crambinae, tribe Haimbachiini. Some authors have placed it in the snout moth The Pyralidae, commonly called pyralid moths, snout moths or grass moths, are a family of Lepidop ...
'' Hübner, 1818 * ''
Yoshiyasua ''Yoshiyasua'' is a grass moth genus (family Crambidae) of subfamily Musotiminae. Some authors have placed it in the snout moth family (Pyralidae), where all grass moths were once also included, but this seems to be in error. The genus contains ...
'' (formerly ''
Melanochroa ''Yoshiyasua'' is a grass moth genus (family Crambidae) of subfamily Musotiminae. Some authors have placed it in the snout moth family (Pyralidae), where all grass moths were once also included, but this seems to be in error. The genus contains o ...
'' Yohiyasu, 1985 ''nec'' Roeder, 1886:
preoccupied The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linn ...
)


References


External links

* Solis, M. Alma (2007)
"Phylogenetic studies and modern classification of the Pyraloidea (Lepidoptera)"
''Revista Colombiana de Entomología''. 33 (1): 1–8. * {{Taxonbar , from=Q248425 Pyralidae Moth families