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Taxonomy
Pyralidae
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The Pyralidae or snout moths are a family of Lepidoptera in the ditrysian superfamily Pyraloidea. In many (particularly older) classifications, the grass moths (Crambidae) are included in the Pyralidae as a subfamily, making the combined group one of the largest families in the Lepidoptera. The latest review by Munroe & Solis, in Kristensen (1999)[full citation needed] retains the Crambidae as a full family of pyraloidea.
Relationship with humans[edit]
Most of these small moths are inconspicuous and of no particular significance to humans. Some are more notable, however. Perhaps the most familiar are waxworms, which are the caterpillar larvae of the greater (Galleria mellonella) and lesser (Achroia grisella) wax moths (subfamily Galleriinae). They are natively pests of beehives, but are bred indoors in enormous numbers as live food for small reptile and bird pets and similar animals. They are also used as fishing bait for trout fishing.
Other notable snout moths are primarily relevant due to their larval food choices. Examples include:
- Alligatorweed stem borer (Arcola malloi: Phycitinae) – biological control of alligatorweed (Alternanthera philoxeroides)
- Almond moth (Cadra cautella: Phycitinae) – pest of stored cereals and dry fruit; now introduced almost world-wide
- Cacao moth, tobacco moth, warehouse moth (Ephestia elutella: Phycitinae) – pest of stored dry vegetable products; Europe, introduced to some other regions (e.g. Australia)
- Dried fruit moth (Cadra calidella: Phycitinae)
- Locust bean moth (Ectomyelois ceratoniae: Phycitinae)
- Etiella behrii (Phycitinae) – pest of stored legumes; Southeast Asia and Australia
- "Flour moths" – pests of stored grains, spices, flour, and similar dry vegetable products; now introduced almost world-wide
- Indian mealmoth (Plodia interpunctella: Phycitinae)
- Mediterranean flour moth, Indian flour moth (Ephestia kuehniella: Phycitinae)
- Grease moth (Aglossa pinguinalis: Pyralinae) – pest of suet and other oily food
- Lesser cornstalk borer (Elasmopalpus lignosellus: Phycitinae) – stalk pest of corn (Zea mays); tropical and subtropical Americas, introduced to the Hawaiian Islands
- Mahogany webworm (Macalla thyrsisalis: Epipaschiinae) – defoliator pest of mahogany trees (Swietenia); Neotropics
- Meal moth (Pyralis farinalis: Pyralinae) – pest of stored grain, flour and other cereals; now introduced almost world-wide
- Pear fruit borer (Pempelia heringii: Phycitinae) – pest of apple and pear fruits; East Asia, introduced to the Hawaiian Islands
- Pine webworm (Pococera robustella: Epipaschiinae) – defoliator pest of pines (Pinus); North America east of Great Lakes region
- Raisin moth (Cadra figulilella: Phycitinae) – pests of stored dry fruit; now introduced almost world-wide
- Rice moth (Corcyra cephalonica: Galleriinae) – pest of stored grain, flour and other cereals
- South American cactus moth (Cactoblastis cactorum: 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: Galleriinae) – pest of stored nuts and drupes; Southeast Asia, introduced to Western Europe
- Sunflower moth (Homoeosoma nebulella: Phycitinae) – pest of sunflower seeds; Europe and surrounding regions
The European corn borer (Ostrinia nubilalis) and southern cornstalk borer (Diatraea crambidoides), formerly considered snout moths, are placed in the Crambidae which, as noted above, are usually regarded as a separate family today.
Systematics[edit]
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 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 moths which are dependent on sloths for their entire life cycle. Most Chrysauginae larvae have a sclerotised ring around seta SD1 of the metathorax.
- Galleriinae (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 (including Endotrichinae, Hypotiinae) – rather diverse in the Old World; a lesser amount of the c.900 species occurs elsewhere. The females of almost all Pyralinae except Cardamyla and Embryoglossa are recognizable by the very short ductus bursae of their genitals.
- Epipaschiinae (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 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.
Genera incertae sedis[edit]
In addition to those assigned to the tribes above, there are several genera of (presumed) Pyralidae which are not firmly placed in this arrangement. 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. Some may even belong to more ancient moth lineages, such as the Alucitoidea or Pterophoroidea. Finally, it is possible that some of these (usually little-studied) genera are junior synonyms of genera described earlier. The genera in question are:
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The following 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 and Tanaobela):
- Alphacrambus
- Peniculimius
- Steneromene
- Thopeutis
- Yoshiyasua (formerly Melanochroa Yohiyasu, 1985 nec Roeder, 1886: preoccupied)
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Diversity description:
There are currently 5 described subfamilies of Pyralidae: Chrysauginae Lederer (1863), Epipaschiinae Meyrick (1884), Galleriinae Zeller (1848), Phycitinae Zeller (1839), and Pyralinae Latreille (1809). Currently there are around 4400 named species of Pyralidae, although much of the diversity is undescribed.
Geographic Range:
Nearctic, Palearctic, Oriental, Ethiopian, Australian, Oceanic Island
Number of stemmata:
from 1 to 6
Secondary setae:
absent
Body setae on verrucae:
absent
Body setae on chalazae:
absent
Body setae on scoli:
absent
Pairs of thoracic legs:
from 3
Larval abdomen description:
A8 SD1 surrounded by chitinous ring (occasionally reduced). A1-A7: L2 anterodorsal or sometimes dorsal of L1. A8 SV-group usually bisetose. A9 L group usually trisetose (L3 rarely lost). Anal shield: distance D1-D1 usually greater than SD1-SD1 (exceptions in Galleriinae, Pyralinae, Euzophera). Crochets in complete circle.
Abdominal glands:
absent
Abdominal prolegs:
present
Pairs of abdominal legs:
from 5
Proleg configuration:
normal
Anal comb on A10:
absent
Reproductive system:
Ditrysian
Oviscapt (ovipositor):
non-piercing
Female genitalia description:
Ostium bursa opening venterally behind sternite 7; ductus bursa generally narrow duct, may be membranous, sclerotized or scobinate; antrum is sclerotized caudal region of ductus bursa; corpus bursa enlarged and sac like, may contain variously shaped signa; accesory sac may arise from corpus bursae; ductus seminalis originates from ductus bursa or corpus bursa; oviscapt is telescopic and is formed from abdominal segments 8 and 9, extension and retraction of oviscapt controlled by muscles inserting on apophyses anteriores (tergite 8) and apophyses posteriores (tergite 9); anal papillae are external membranous setose lobes formed from segments 9-10
Female pregenital sexual scales:
absent
Female accessory glands:
one pair
Female oviduct opening:
below anus
Female bursa ostium opening:
between S7 and venter 8
Female anterior apophyses originating:
originating from T8
Male coremata:
absent
Male pregenital sexual scales:
present, absent
Male genitalia description:
tegumen and vinculum form complete ring; vinculum elongated anteriorally into saccus; uncus usually well developed, although may be reduced; uncus with ventro-anteriorly elongate arms articulating with gnathos; uncus may be setose caudally; gnathos variable, may be present or reduced; distal end of gnathos variously modified; valves often simple, although may be lobed or setose; transtilla variably developed; juxta platelike; aedoeagus (phallotheca) short cylinder; caecum present or absent; cornuti present or absent
Sternum 5:
without fenestra
Sternum 5 gland:
absent
Male has:
phallotheca and aedeagus (phallus)
Epiphysis:
present
Adult thorax description:
Males may have androconia.
Forelegs:
normal, modified
Leg description:
Variable. From long and slender to short and stout. May be smoothly scaled or with modified scales. Some males with prominent androconia.
Wing venation??description:
Forewing of variable shape. Forewing veins with R2 closely apposed to R3 and R4, and not usually stalked; R3, R4 and R5 may be reduced to one or two veins; M1 originates near anterior angle of discal cell; M2, M3 and CuA1 originate from posterior angle of cell; M2 and M3 may be stalked; CuP well developed, incomplete, or absent and reduced to a fold; 1A strongly developed; 2A distally free or connected to 1A by a crossvein to form a closed cell, sometimes with a free portion extending beyond this cell. Hindwing wide, with narrow fringe. Hindwing veins Sc+R1 and Rs may be anastomosed or separate; M2 and M3 usually separate, but may be fused; CuA1 and CuA2 usually arising separately from discal cell; CuA1 free, rarely fused with M3; CuP and 1A+2A present.
Wing venation:
heteroneurous
Forewing basal loop:
absent
Forewing pterostigma:
absent
Forewing chorda:
absent
Forewing upper surface with microtrichia:
absent
Hindwing anal vein notation:
In Phycitinae, radial sector free or partially fused with Sc+Rs. M2+3 fused (partially or fully), or separate. Cell often open. CuP absent. All three Anal Veins generally present.
Hindwing pterostigma:
absent
Wing coupling:
present, with frenulum
Forewing description:
Both sexes with a retinaculum of stiff scales on underside of cubital area. Males may have sclerotized frenulum hook, but is lost in some groups.
Hindwing description:
Frenulum single in males, and may be single or multiple in females.
Abdomen tympanum:
present
Abdomen tympanum description:
Tympanal cases closed; conjunctiva and tympanum in the same plane; praecinctorium absent; secondary venulae present or absent
Thorax tympanum:
absent
Ocelli:
present, absent
Eyes:
smooth
Labial palpus:
porrect, upcurved
Labial palpus modification:
Labial palps have also been described as obliquely ascending
Maxillary palpus:
present, absent, minute
Proboscis:
present, absent, reduced
Proboscis texture:
scaled
Proboscis description:
Densely scaled on exposed basal anterior surface
Mandibles:
absent, reduced
Head vertex scaling:
normal
Female antennae:
bipectinate, filiform, pectinate
Male antennae:
bipectinate, filiform, pectinate, thickened
Male pedicel description:
In Phycitinae the scape may posses various modification, such as having a long spine or being deeply notched.
Male flagellomere description:
In Phycitinae the basal flagellomere may posses various modifications, mainly in the form of a high concentration of scale-like sensilla.
Antennal sensillum:
Antennal sensillum present
Basiconium:
few
Sensillum vesiculocladum:
absent
Asciod sensilla:
absent
General antennae description:
Often sexually dimorphic
Adult head description:
Frons rounded, scaling smooth; chaetosema often present, but reduced, forms a radiating group of fine seta near posteroventral angle of vertex
Apomorphies:
Forewing vein R5 stalked or fused with R3+R4. Tympanal case closed, or nearly closed. Praecinctorium absent. Tympanum and conjunctiva lie in the same plane. Male genitalia with uncus arms, a pair of processes arising laterally from the base of the uncus. Larvae almost always with sclerotized ring around base of seta SD1
Animal / pathogen
Entomophthora neopyralidarum infects adult of Pyralidae
Animal / parasitoid / endoparasitoid
larva of Eurysthaea scutellaris is endoparasitoid of larva of Pyralidae
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