AMPHIESMENOPTERA - the “garment wings” The Major Groups of Holometabola Holometabola (=Endopterygota) Order Hymenoptera Superorder Neuropterida Order Raphidioptera Order Megaloptera Order Neuroptera Order Coleoptera Order Strepsiptera, incertae sedis Superorder Mecopterida (= Panorpida) Antliophora Order Mecoptera - scorpionflies Order Siphonaptera - fleas Order Diptera - true flies Amphiesmenoptera Order Trichoptera - caddisflies Order Lepidoptera - butterflies & moths 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 Biodiversity in Insects: 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 taxonomy: 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 animal matter • Many are of great economic importance LEPIDOPTERA - butterflies and moths Diversity and Distribution • 150,000 described species 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” “mandibulate Micropterygidae Agathiphagidae 1 Heterobathmiidae Eriocraniidae 2 Acanthopteroctetidae GLOSSATA 3 Lophocoronidae COELOLEPIDA 4 Neopseustidae 5 EXOPORIA 7 6 (Mnesarchaeidae+Hepialoidea) MYOGLOSSATA HETERONEURA NEOLEPIDOPTERA LEPIDOPTERA - butterflies and moths Synapomorphies of the major groups 1- Agathiphagidae + Heterobathmiidae + GLOSSATA • Adult paraglossae lost • Pupal mandible hypertrophied, angularly bent • Larva with only one maxillary endite lobe (probably the galea) • Larva with metathoracic spiracle non-functional 2 - GLOSSATA • Mandible non-functional after pupa/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” (Nepticuloidea, Incurvarioidea, etc.) Tineoidea 8 9 Gracillarioidea Yponomeutoidea DITRYSIA Gelechoidea Zygaenoidea + Sesioidea/Cossoidea Tortricoidae 10 Pyraloidea 11 APODITRYSIA Noctuoidea OBTECTOMERA 12 Bombycoidea RHOPALOCERA MACROLEPIDOPTERA Drepanoidea Geometroidea 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 - OBTECTOMERA • 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 + Papilionoidea 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) • Basal 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 “caterpillars” 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 Insect of Australia, 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 genus, Agathiphaga 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, Agathis (Araucariaceae). Common name "kauri moths". LEPIDOPTERA - butterflies and moths Synopsis of non-Heteroneuran Lepidoptera Heterobathmiidae • Recently discovered genus Heterobathmia of southern South America with only 10 species • Small, diurnal moths resembling Eriocraniidae • Found on flowering Nothofagus 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