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Home , Acanthopteroctetidae, Agathiphaga, Amphiesmenoptera, Coelolepida, Ditrysia, Eriocraniidae

AMPHIESMENOPTERA - the “garment wings” The Major Groups of Holometabola Holometabola (=) Superorder Order Raphidioptera Order Order Order Coleoptera Order , Superorder Mecopterida (= ) Antliophora Order - scorpionflies Order Siphonaptera - Order Diptera - true Order Trichoptera - Order - & AMPHIESMENOPTERA - the “garment wings”

SYNAPOMORPHIES • Adult prelabium fused with hypopharynx • Pronotum with paired setose “warts” • Wings with extensive covering of setae • Anal veins of the forewing apparently looping up into a double-Y configuration • Presence of paired glands opening on sternum V • Male genital segment IX with tergum and sternum fused, forming a closed ring AMPHIESMENOPTERA - the “garment wings”

SYNAPOMORPHIES (continued)

• Anterior margin of female segment VIII and IX with long, rodlike apodemes accommodating the insertion of protractor/retractor muscles on the extensible oviscapt • Female heterogamety (female is XY, male is XX) • Chromosome number specialized (high, n=30 in Trichoptera, n=31 in Lepidoptera) • Larval prelabium and hypopharynx fused into a lobe apically carrying the salivary (silk) gland orifice

And many more (see Kristensen, N.P. 1984. Studies on the morphology and systematics of primitive Lepidoptera. Steenstrupia 10(5): 141-191; Kristensen, N.P. 1997. Early evolution of the Lepidoptera + Trichoptera lineage: phylogeny and the ecological scenario. In: Grandcolas, P. (editor). The Origin of in : Phylogenetic Tests of Evolutionary Scenarios. Mém. Mus. Natn. Hist. Nat., 173: 253-271.) Hypothetical amphiesmenopteran, from Kristensen (1984) LEPIDOPTERA - butterflies and moths LEPIDOPTERA - butterflies and moths SYNAPOMORPHIES • Wings with dense covering of broad scales • Wings with M 3-branched • Foretibia with single apical spur (or absent) • Foretibia with “epiphysis” on inner surface, spurlike process used to clean antennae • Median ocellus lost • Cerci lacking in both sexes • Tergum I desclerotized And many more (see Kristensen, N.P. 1984. Studies on the morphology and systematics of primitive Lepidoptera. Steenstrupia 10(5): 141-191. See also references cited therein) See also the most important work published to date on Lepidoptera phylogeny and : N.P. Kristensen (editor). 1999. Lepidoptera, Moths and Butterflies, volumes 1 & 2. Handbook of Zoology, vol. 4, part 35. Walter de Gruyter Publishers. Berlin. LEPIDOPTERA - butterflies and moths Habitat & Habits

• Adult moths generally nocturnal, butterflies and skippers diurnal • Adults suck nectar or feed on pollen; a very few noctuids suck blood • Largest group of plant feeding organisms, probably the most recent to have radiated with the angiosperms • Most are external leaf feeders, but habits and habitats of the larvae include leaf, woody stem, and fruit and nut borers; stored products and detritus feeders; fungus and lichen feeders; a few parasites and predators; some feed on decomposing plant and matter • Many are of great economic importance LEPIDOPTERA - butterflies and moths Diversity and Distribution • 150,000 described worldwide, but probably 300,000 - 500,000 in existence, in 46 superfamilies and 121 families • About 12,000 species in U.S. and Canada “mandibulate moths”

Micropterygidae

Agathiphagidae 1 Heterobathmiidae

Eriocraniidae 2

GLOSSATA 3 Lophocoronidae 4 5 7 6 (Mnesarchaeidae+) LEPIDOPTERA - butterflies and moths Synapomorphies of the major groups 1- Agathiphagidae + Heterobathmiidae + • Adult paraglossae lost • Pupal mandible hypertrophied, angularly bent • with only one maxillary endite lobe (probably the galea) • Larva with metathoracic spiracle non-functional 2 - GLOSSATA • Mandible non-functional after /adult ecdysis, with genuine articulation basally (and often strongly reduced) • Maxilla devoid of sclerotized lacinia • Maxillary galeae forming a proboscis, spirally coiled in repose • Larva with articulated “spinneret” on the apex of the prelabio-hypopharyngeal lobe 3 - COELOLEPIDA • Hollow scales on wing surface 4 - Lophocoronidae + MYOGLOSSATA • Vein R (versus Rs) always unforked 5 - MYOGLOSSATA • Proboscis with an intrinsic musculature 6 - NEOLEPIDOPTERA • Pupa adecticious [pupa with immobile mandibles] • Pupa obtect [wings and appendages are appressed to the body and most abdominal segments are immovable], with backward pointing spines dorsally • Larva with musculated, crochet-bearing prolegs on abdominal segments III-VI & X 7 - EXOPORIA • Female genitalia exoporian HETERONEURA “monotrysian Heteroneura” (, Incurvarioidea, etc.)

Tineoidea 8 9 Gelechoidea

Zygaenoidea + /

Tortricoidae 10 11 APODITRYSIA

OBTECTOMERA 12 RHOPALOCERA LEPIDOPTERA - butterflies and moths Synapomorphies of the major groups

8- HETERONEURA • Venation heteroneurous due to simplification of hind wing venation (Sc and R fused and Rs unbranched) • Wing coupling with retinaculum and frenulum and jugum reduced in males 9 - DITRYSIA • Internal connection between copulatory opening and ovipore - ditrysian type • Large apodemes on abdominal sternite 2 • Muscles in proboscis with short bands, not long fibers 10 - APODITRYSIA • Apodemes of abdominal sternum 2 specialized, with short, large bases 11 - • Pupal abdominal segment 1-4 immobile • Dorsal row of spines lost on pupal tergites 12 - MACROLEPIDOPTERA • Complete loss of CuP • Larval proleg crochets in a “mesoseries” LEPIDOPTERA - butterflies and moths Synapomorphies of the major groups

13 - RHOPALOCERA • Abdominal tergite 1 pouched Hedylidae • Eggs resting upright • Pupa girdled • 7 other characters 13 14 - Hesperiidae + Hesperiidae • Brain with large optic lobe • 5 other characters RHOPALOCERA 14 15 - Papilionoidea 15 • 5 technical adult thoracic Papilionoidea characters LEPIDOPTERA - butterflies and moths Major features in lepidopteran evolution (more or less chronologically)

• families mandibulate, without proboscis • Proboscis (elongate, fused, “zipped” galeae of maxillae) evolved in Glossata; adult mandibles non-function or vestigial; proboscis extended by hemolymph with aid of resilin filament • Glossatan larvae with silk-secreting spinneret at tip of fused prelabium and hypopharynx • Hollow scales evolved in the Coelolepida; abwing surface with windows and perforations, also embedded with pigment granules • Adectitious, obtect pupae occurred in the Neolepidotera, • Double layer of wing scales evolved in the Neolepidoptera LEPIDOPTERA - butterflies and moths Major features in lepidopteran evolution (more or less chronologically) • Typical neolepidopteran larvae or “” have musculated abdominal prolegs with crochets on segments 3-6 and 10 • Female neolepidopteran with separate copulatory and oviposition openings, but with no connection between the two (monotrysian condition, with a common cloaca opening to the outside is the primitive condition) • Retinaculo-frenate wing coupling occurred in the Heteroneura (jugate wing coupling the primitive condition) • In Ditrysia an internal connection evolved between the separate copulatory and oviposition openings - dytrisian condition • Basal ditrysian caterpillars are leaf miners, but most others external leaf feeds, some in webs or cases LEPIDOPTERA - butterflies and moths Mouthparts LEPIDOPTERA - butterflies and moths Wing coupling

From of , 2nd edition LEPIDOPTERA - butterflies and moths Female reproductive systems - monotrysian LEPIDOPTERA - butterflies and moths Female reproductive systems - exoporian LEPIDOPTERA - butterflies and moths Female reproductive systems - ditrysian LEPIDOPTERA - butterflies and moths Larva and pupa LEPIDOPTERA - butterflies and moths Synopsis of non-Heteroneuran Lepidoptera Micropterygidae - 120 species • Small, about 1 cm, generally diurnal moths with metallic-colored wings • Adults eat pollen, some feed on fern spores • Larvae feed on fungal hyphae, bryophytes, or lower parts of grasses, other flowering plants, or decaying material • Living sister group to all other Lepidoptera LEPIDOPTERA - butterflies and moths Synopsis of non-Heteroneuran Lepidoptera

Agathiphagidae • One , with 2 species, from southwest Pacific region • Adults nocturnal, small to medium sized, and superficially very similar to caddisflies • Mandibles are large, but it is unknown if they are functional • Larvae are apodous miners in seeds of kauri pines, (Araucariaceae). Common name "kauri moths". LEPIDOPTERA - butterflies and moths Synopsis of non-Heteroneuran Lepidoptera

Heterobathmiidae • Recently discovered genus of southern South America with only 10 species • Small, diurnal moths resembling • Found on flowering where adults eat pollen • Larvae are leaf miners in Nothofagus; larvae can leave and enter a new leaf, this behavior not common among other leaf mining insects LEPIDOPTERA - butterflies and moths Synopsis of non-Heteroneuran Lepidoptera Eriocraniidae Exclusively Holarctic family of about 20 described species. Small, diurnal moths with iridescent wing patterns. Do not visit flowers, but use short proboscis for drinking water. Mandibles are present, but nonfunctional. Larvae are apodous leaf miners, almost exclusively of trees in orders and Lophocoronidae Small Australian genus Lophocorona of 6 species. Small, nocturnal moths. Females and immatures unknown Acanthopteroctetidae Small western Nearctic genus Acanthopteroctetes with 4 species. Predominantly diurnal moths. Known larvae are leaf miners. LEPIDOPTERA - butterflies and moths Synopsis of non-Heteroneuran Lepidoptera

Neopseustidae • About 12 described species from southern Asia and South America • Medium-sized, generally nocturnal moths • Immature stages remain unknown Mnesarchaeidae • Single genus with 14 described species from New Zealand • Small, diurnal moths with well developed proboscis used to drink water, but have not been observed at flowers • Larvae are ground dwelling and feed on leaf litter detritus LEPIDOPTERA - butterflies and moths Synopsis of non-Heteroneuran Lepidoptera

Hepialoidea - 600 species • Ghost moths and relatives; some very large • Primarily restricted to southern continents • Larvae tunnel in woody tissue above and below ground or feed on roots externally, some also feed on liverworts or fungi; many construct silk galleries in the litter or below ground

Palaeosetidae - Australian/Oriental/S. America Neotheoridae - S. American Anomosetidae - Australian Prototheoridae - S. African - cosmopolitan Hepialidae LEPIDOPTERA - butterflies and moths Diversity and Distribution (selected families) Heteroneura - Nepticuloidea - 800 species • Very small moths, 1.5-5 mm; adults run and erratically • Larvae are leaf miners, but some found in twigs, fruit, or produce tiny galls; occur on many plant families • Very seldom collected, probably many new species (Andesianidae - 3 species) • Southern Argentina and Chile • New family described only in 2003! LEPIDOPTERA - butterflies and moths Diversity and Distribution (selected families) Heteroneura - Incurvarioidea - 90 species, mostly North America • Larvae bore in seeds, fruits, or stems of many plant families • Includes several genera and species of Yucca moths, involved in complex symbiotic associations with Yucca (Agavaceae) (see the work of Pellmyr) • Female punctures pistil of yucca flowers and inserts 1 egg. Then packs pistil with pollen which ensures pollination and a subsequent seed supply for larva • Females have curved maxillary “tentacles” used to collect yucca pollen - fairy moths; 300 species • Small moths with very long, filiform antennae - easily mistaken for leptocerid and macronematine caddisflies • Little known of their biology

Incurvariidae - 100 species • Small, dark moths with folded maxillary palps • Larvae begin as leaf miners, then exit and construct an oval case of two leaf pieces • Case construction and continued feeding results in oval holes in the leaf Prodoxidae Adelidae

Incurvariidae LEPIDOPTERA - butterflies and moths Diversity and Distribution (selected families) Heteroneura - - 3,000 species • Larvae construct cases of frass and the material they feed on • Include species that feed on plant and animal detritus, dung, lichen, bracket fungi, grains, feathers, keratin, and detritus in animal nests • Several species feed on wool clothing, rugs, etc - clothes moths • On many occasions people have sent me tineid larvae they were convinced were remarkable terrestrial larvae, only to be disappointed LEPIDOPTERA - butterflies and moths Diversity and Distribution (selected families) Heteroneura - Tineoidea Psychidae - bagworms; 1,000 species • “Bagworms” build portable, spindle-shaped cases of leaf, needles, or twigs from their host plant • Females usually apterous or brachypterous or neotenic and remain in case where they mate with males; attract male with pheromones; he mates with female through opening at end of case • Mouthparts vestigial in both sexes and adults do not feed • Some females parthenogenetic LEPIDOPTERA - butterflies and moths Diversity and Distribution (selected families) Heteroneura - Gracillarioidea - leaf blotch miners; 2,000 species • Small to minute moths with lanceolate wings; adults sit with anterior end of body elevated and wing tips touching the surface • Larvae are the “leaf blotch miners” and the most common species in North America mine in white and aspen; usually pupate in the mine LEPIDOPTERA - butterflies and moths Diversity and Distribution (selected families) Heteroneura - Yponomeutoidea Yponomeutidae - ermine moths, ca. 500 species • Small, brightly colored moths, or white and silvery. • Most are external leaf feeders as larvae, but also includes those that live in frail silken communal webs (ailanthus webworm) and pine needle sheath miner as well as those that bore in twigs, buds, fruits, or are leaf miners • Adults of some species rest with the head on the substrate and the body up LEPIDOPTERA - butterflies and moths Diversity and Distribution (selected families) Heteroneura - Yponomeutoidea - diamondback moths, ca. 50 species • Most distinctive because the males of some species have diamond shaped markings on the wings when they are folded • Larvae feed principally on Brassicaceae • One species, xylostella, is a pest of cabbage; larvae live in loose webs and skeletonize the leaves; this species is also known for its long migrations LEPIDOPTERA - butterflies and moths Diversity and Distribution (selected families) Heteroneura - - ca. 3000 species • Adults are small moths, the labial palps are long and upcurved and the hind wing is trapezoidal in shape with a tapered apex • Larval habits are varied, include some leaf miners, leaf tiers and rollers, a few gall formers, especially in golden rod, and a pest of stored grains, the Angoumois grain , Sitotroga cerealella as well as the pink boll worm of cotton and a pest of potatoes, thus the family is of great economic importance LEPIDOPTERA - butterflies and moths Diversity and Distribution (selected families) Heteroneura - Gelechioidea Oechophoridae - ca. 3000 species • Adults are often brightly colored, broad, squarish moths with upturned labial palps • Larvae of some species feed on highly toxic plants such as cow parsnip • Some larvae are predators of scale insects and aphids • Others feed in webs, rolled or tied leaves, on dead leaves, on decaying fungi, or even animal carcasses