The Pyraloidea are one of the larger superfamiles of the Lepidoptera. The two families, Pyralidae and Crambidae, comprise about 16,000 named species. Many species remain undescribed.
The snout moths (Pyraloidea) contain over 16,000 described species worldwide (Solis, 1997, Munroe & Solis 1998). The life histories of pyraloid moths are very diverse. Some caterpillars feed on living plant tissue (as stem borers, leaf, root, or seed feeders), while others are predators or live parasitically in the nests of ants, bees, and wasps. Many species specialize in the exploitation of dried or decaying plant or animal substrate, and several are significant pests of important crop species and stored products (Solis 1996).
Nearctic, Palearctic, Oriental, Ethiopian, Neotropical, Australian, Oceanic Island
Geographic Range description:
Pyraloidea occur on all continents except the Antartica, and on most islands, including oceanic ones such as Hawaii, Rapa, the Azores, St. Helena and Amsterdam Island. Though best respresented at low and middle elevations in the tropics, tehy range from the high arctic of Ellesmere Island to Tierra del Fuego and from equatorial lowlands to alpine snowline. Their patterns of distribution indicated repeated dispersal at many evolutionary levels. Some nearly cosmopolitan species have been spread by human agency. In numbers of species the Oreiental and Neotropical faunas are almost the same, and the Australian fauna about three-fourths as large. The smallest pyraloid fauna is the Nearctic; the Ethiopian is not much larger, and the Palearctic is almost as large as teh two together.
Adult Abdomen Morphology
Female genitalia description:
Primitively with a pair of membranous setose ovipositor lobes, approximated or fused dorsally, that of each side supported by usually T-shaped posterior apophysis. Ostium base simple or variously armed. Corpus bursae variable in shape and armature; ductus bursae usually differentiated from corpus bursae; sometimes an accessory sac arising from corpus bursae; ductus seminalis usually arising from ductus bursae, but sometimes from corpus bursae; ductus bursae often with a collarlike or complex sclerite at or just posterior at opening of ductus seminalis.
Male genitalia description:
Tegumen and vinculum forming a complete ring, often with compelx lateral folding; vinculum usually produced anteriad into a midventral saccus. Uncus various, almost always present. Gnathos primitively consisting of a pair of lateral arms arising from junction of tegumen and uncus, medially fusing to form a posteriorly directed median process. Basal articulations of gnathos either movable,hinging at tegumino-uncal suture, or fixed, in the latter case often joined to distolateral margins of tegumen. Transtilla a transverse sclerotized band between costae of valvae, often medially narrowed or absent, rarely with special armature. Phallus a variously sclerotized tube, sometimes distorted or ornate, projecting through a membranous or spinulose sleeve, the manica, or sometimes a sclerotized troughlike or tubular anellus. Inside phallus a membranous eversible vesica, generally minutely spinulose, often also with single or grouped larger spines, the cornuti; the latter sometimes deciduous, capable of being shed into the female bursa during copulation. Juxta variable in shape and size, ventral to manica or anellus, connecting ventrolaterally with bases of valvae. Valva extremely various in form, primitively with subparallel costal and ventral margins, each somewhat inflated, the latter inflation known as the sacculus, and with rounded or oblique terminal margin; costa articulating basally with transtilla, most of the base hinged with vinculum; ventral angles articulating with juxta; mesal surface generally with variously grouped setae and sometimes spines, often also with variously disposed ridges and processes; in certain groups the costa and/or sacculus produced distally as free process.
Adult Thorax Morphology
Adult thorax description:
In males often with variously disposed androconia; thoracic structure variable, but not studied comparatively. Wings variable in shape; forewing wide to narrow, hindwing wide, with fringe much narrower than membrane; costa usually straight or arched, but sometimes concave, sinous or distorted in either forewing or hindwing, especially in males;termen usually convex or sinuate, tending to be more strongly curved in cell M3.
Number of tibial spurs foreleg:
Number of tibial spurs midleg:
Number of tibial spurs hindleg:
Slender to thick and robust, smoothly scales or with tcick vestiture of normal or modified scales, in males often withprominent androconia, sometime in retractile tufts or with accompanying distortions of the leg.
Forewing primitively with Sc, 5-branched R, R3 and R4 stalked, no areole, 3-branched M, M2 closer to M3 than to M1, stem of M absent or faintly indicated, discal cell usually closed, CuA 2-branched, CuP represented by a vein, fold, a weak tubualar vestige adjacent to termen, or absent. 1A strong and tubular, 2A shorter and weak,often joined to 1A by a crossvein. Hindwing with Sc+R1 approaching or anastomosed with Rs for some distance beyond end of discal cell, then diverging again, Rs not branched, basally often short-stalked with M1 from anterior angle of cell, but sometimes connate with M2 or the later arising separately form discocellular, M 3-branched, M2 closer to M3 than to M1, as on forewing, CuA 2-branched, CuP usually well developed, 1A+2A always and 3A usually present. Discal cell open in many Crambinae. Stem of CuA in some groups with pecten of fine setae.
Forewing cell veins:
Number of Rs veins in forewing:
Number of M veins in forewing:
Forewing upper surface with microtrichia:
Number of Rs veins in hindwing:
Number of M veins in hindwing:
Wing coupling description:
Both sexes with a retinaculum of stiff scales on underside in cubital area, males primitively with a sclerotized frenulum hook extending posterodistad from behind costa, but the frenulum hook lost in some groups. Frenulum unisetose in male, multi- or uni-setose in female.
Abdomen tympanum description:
Tympanal organ with paired tympanal chambers on the ventral part of the base of the abdomen, facing more or less forward into space between the thorax and abdomen; rarely reduced or absent. Tympanal case either almost completely closed (Pyralidae) or open, with a wide anteromedial aperture (Crambidae). Conjunctiva and tympanum in the same plane (Pyralidae) or not in the same plane, meeting at a distinct angle (Crambidae). Praecinctorium absent (Pyralidae) or present (Crambidae).
Adult Head Morphology
porrect, upcurved, large
Number of labial palp segments:
Labial palpus modification:
normally prominent, articulations with limited movement, rarely reduced, sometimes excavated dorsally to receive maxillary palpus; normal position decurved, porrect, obliquely ascending, or upturned against frons or even over vertix; scaling well-developed, various, sometimes including androconia.
Number of maxillary palp segments:
from 1 to 4
Number of chaetosomata:
present, absent, reduced
primitively well-developed, coiled when not in use, exposed anterior surface of basal coil densely scaled; sometimes reduced or absent; even very rudimentary proboscides often retain basal scaling.
Head vertex scaling:
General antennae description:
Various, filiform, annulate, prismatic, laminate, lesss commonly uni- or bi-pectinnate; often sexually dimorphic
Adult head description:
Frons smoothly or roughly scaled; rounded, flat and oblique, or less commonly, prominent or with variously shaped spines, ridges or processes. Vertex with erect scaling in tufts. Maxillary palpus with scaling compressed , distally dilated , plumose, or forming an aigrettelike tuft. Ocellus most often present, situated near dorsal margin of eye, reduced or absent in some species and genera. Chaetosema often present, but in reduced form, as a somewhat radiating group of short fine setae near posteroventral angle of vertex; a raised boss or specialized setal bases rarely visible on the denuded vertex.
with paired abdominal tympanal organs consisting of paired tympanal chambers on the ventral part of segment two, supporting a tympanum and conjunctiva, the former with a sensory scoloparium attached; maxillary palpus usually present; proboscis, if present, scaled at the base; R3 and R4 of the forewing stalked or fused; Sc+R1 and Rs of the hindwing are anastomosed or approximated for some distance distad of the discal cell
Life History and Behavior
Pyraloid larvae are characteristically concealed feeders: folding, rolling, webbing or tying leaves; making sand, silk or frass tunnels or tubes; boring in stems, roots, shoots, buds, fruits, or galls; mining leaves; making cases of various kinds; or sheltering in nests or ants, bees, wasps and termites.
Life History: Immature Stages
The great majority of present-day pyraloid larvae feed on the tissues of vascular plants, either gymnosperms or angiosperms. In addition to plant-feeders and to scavengers, there are some species predaceous on Homoptera and other insects,or that are inquilines or parasites in the nests of social insects.
Evolution and Systematics
Systematic and taxonomic history
The definition of the Pyraloidea has narrowed considerably over the past few decades; classically the Pyraloidea included Pterophoridae, Alucitidae, Thyrididae, Hyblaeidae, Oxychirotidae, and Tineodidae (reviews in Fletcher & Nye 1984; Minet 1986; Nielsen 1989; common 1990). Minet (1986) included the Dudgeoneidae, but later withdrew this placement (Minet 1991). With subsequent work all families were transferred to other superfamilies or to their own superfamily leaving only the Pyralidae (sensu lato). Munroe (1972) separated it into 3 informal groups based on the tympanal organ: Pyraliformes, Crambiformes, and Midiliformes. Minet (1983), in his broad study of tympanal organs, showed that those of the Midilinae were of the crambiform type, though reduced. He separated the Pyralidae (sensu lato) into two sister families, Pyralidae (sensu stricto), corresponding to Pyraliformes, and Crambidae, comprising Crambiformes plus Midiliformes of Munroe (1972). Munroe & Solis (1999) followed this classification because the morphological and cladistic evidence for the monophyly of the Pyralidae and Crambidae and their sister group relationship is clear enough to warrant their separation at the family level. The sister groups to the Pyraloidea have been hypothesized to be the Thyridoidea and Hyblaedoidea, but no comparative morphological study has been conducted.
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