AN INTRODUCTION TO MICROLEPIDOPTERA: Phylogeny, Classification, and Morphology (gag me with a spoon!) John W. Brown USDA, Systematic Entomology Laboratory U.S. National Museum of Natural History Washington, DC 20013-7012

General Remarks Microlepidoptera – a category of convenience (not all are small). Approximately 40-45% of the order (described, that is). Majority of the fundamental developments characteristic of the order take place within Microlepidoptera. Phylogeny is poorly understood.

Overview of Presentation Briefly summarize classification based on phylogeny from Kristensen (1998). Briefly discuss major lineages and morphological developments that define them. Examine in detail adults of a few of the major lineages (mini-labs).

Kristensen 1998 Handbook of Zoology

Micropterigidae (Zeugloptera) 1 family with 121 described species Tiny moths, 10 mm or less Fuzzy head Metallic colored wings

Micropterigidae (Zeugloptera) 1 family with 121 described species Homoneurous wings Jugate wing coupling

Micropterigidae (Zeugloptera) 1 family with 121 described species Homoneurous wings Jugate wing coupling Monotrysian female reproductive system

Micropterigidae (Zeugloptera) 1 family with 121 described species Homoneurous wings Jugate wing coupling Monotrysian female reproductive system Strongly asymmetrical mandibles, short labial palpi

Eriocraniidae (Glossata) 1 family with 24 described species Tiny moths, 10 mm or less Fuzzy head Metallic colored wings

Eriocraniidae (Glossata) 1 family with 24 described species Homoneurous wings Jugate wing coupling

Eriocraniidae (Glossata) 1 family with 24 described species Homoneurous wings Jugate wing coupling Monotrysian female reproductive system

Eriocraniidae (Glossata) 1 family with 24 described species Homoneurous wings Jugate wing coupling Monotrysian female reproductive system Sucking mouthparts – non-functional mandibles, galea forming proboscis

Hepialoidea (Exoporia) 5 families with ca. 500 described species Wingspan up to 25 cm Adults crepuscular/nocutrnal As many as 30,000 eggs “broadcast” by female in flight

Hepialoidea (Exoporia) 5 families with ca. 500 described species Homoneurous wings Jugate wing coupling

Hepialoidea (Exoporia) 5 families with ca. 500 described species Homoneurous wings Jugate wing coupling Exoporian female reproductive system

Hepialoidea (Exoporia) 5 families with ca. 500 described species Homoneurous wings Jugate wing coupling Exoporian female reproductive system Mouthparts reduced – proboscis absent or short

Incurvarioidea (Heteroneura - Monotrysia) 5 families with ca. 410 described species Heliozelidae, Adelidae, Prodoxidae, Cecidosidae, Incurvariidae Small to tiny moths, forewing length 1.7-16 mm

Incurvarioidea (Heteroneura - Monotrysia) 5 families with ca. 410 described species Heteroneurous wings Frenate wing coupling

Incurvarioidea (Heteroneura - Monotrysia) 5 families with ca. 410 described species Heteroneurous wings Frenate wing coupling Monotrysian female reproductive system

Incurvarioidea (Heteroneura - Ditrysia) 5 families with ca. 410 described species Heteroneurous wings Frenate wing coupling Monotrysian female reproductive system Mouthparts reduced – proboscis absent or short

Incurvariidae – yucca moths

Tineiodea (Heteroneura - Ditrysia) 5 families with ca. 4,200 described species Tineidae, Eriocottidae, Lypusidae, Acrolophidae, Psychidae, Arrhenophanidae The most primitive ditrysians

Tineiodea (Heteroneura - Ditrysia) 5 families with ca. 4,200 described species Heteroneurous wings Frenate wing coupling

Tineiodea (Heteroneura - Ditrysia) 5 families with ca. 4,200 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system

Tineiodea (Heteroneura - Ditrysia) 5 families with ca. 4,200 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system Mouthparts well developed – proboscis reduced

Tineiodea mini-lab Tineidae Erect scales on head Inconspicuous maxillary palpi Short labial palpi Acrolophidae Fuzzy head and body Large, dark adults Males often with elongate palpi Psychidae Wings dark-smoky, scales usually lost Short antennae Characteristic larval cases

Gracillarioidea (Heteroneura - Ditrysia) 4 families with ca. 2,400 described species (mostly Gracillariidae) Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system Mouthparts well developed; labial palpi with lateral bristles (also present in Tineoidea)

Yponomeutoidea (Heteroneura - Ditrysia) 6-9 families with ca. 1,500 described species Classification unstable (duh!): Yponomeutidae, Plutellidae, Ypsolophidae, Acrolepiidae, Glyphipterigidae, Argyresthiidae, Heliodinidae, Lyonetidae (sometimes families, sometimes subfamilies) [Galacticidae] A heterogenous assemblage of relatively primitive micros. Autapomorphy: pleural lobes just before genitalia – posterior expansion of pleuron VIII

Yponomeutoidea (Heteroneura - Ditrysia) 6-9 families with ca. 1,500 described species Heteroneurous wings Frenate wing coupling

Yponomeutoidea (Heteroneura - Ditrysia) 6-9 families with ca. 1,500 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system

Yponomeutoidea (Heteroneura - Ditrysia) 6-9 families with ca. 1,500 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system Mouthparts well developed; labial palpi variable but always large and conspicuous

Yponomeutoidea mini-lab Smooth scaled head Labial palpi variable Proboscis unscaled Apex of hindwing usually rounded; more rarely hindwing oblong-lanceolate with pointed apex Yponomeutidae Yponomeuta Yponomeutidae Atteva Plutellidae Plutella Galacticidae Homadaula

Gelechioidea (Heteroneura - Ditrysia) 10-25 families with ca. 16,500 described species Classification unstable (what a surprise): Gelechiidae, Elachistidae (Stenomatidae, Ethmiidae, Depressariidae, Agonoxenidae), Xyloryctidae (Scythrididae) Schistomeoidae, Oecophoridae (Stathmopodidae), Amphisbatidae, Lecithoceridae, Batrachedridae, Deocloniidae, Coleophoridae (Blastobasidae, Momphidae), Autostichidae (Symmocidae), Cosmopterigidae, others?

Gelechioidea (Heteroneura - Ditrysia) 10-25 families with ca. 16,500 described species Heteroneurous wings Frenate wing coupling

Gelechioidea (Heteroneura - Ditrysia) 10-25 families with ca. 16,500 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system

Gelechioidea (Heteroneura - Ditrysia) 10-25 families with ca. 16,500 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system Mouthparts well developed; labial palpi large, upturned

Gelechioidea mini-lab Ethmiidae Smooth scaled head Long upturned labial palpi Proboscis scaled Large hindwing with rounded apex Characteristic black and white forewing pattern Ethmiidae Ethmia Ethmiidae Ethmia Ethmiidae Ethmia

Gelechioidea mini-lab Coleophoridae Labial palpi usually not as conspicuously upturned Lanceolate wings (pointed apically) Inconspicuous paired patches of special scales/spines subdorsally on most abdominal segments

Gelechioidea mini-lab Oecophoridae Smooth scales on head Large upturned labial palpi Proboscis scaled Apex of hindwing rounded Oecophoridae Promalactis Oecophoridae Callima Oecophoridae Pleurota

Gelechioidea mini-lab Blastobasidae (ugly brown moths) Smooth scales on head Shorter upturned labial palpi Proboscis scaled Lanceolate hindwing Rows of bronze spines on abdominal segments Males of some species with notch near base of antenna Blastobasidae Blastobasis

Gelechioidea mini-lab Stenomatidae Smooth scaled head Long upturned labial palpi Proboscis scaled Large hindwing with rounded apex [Hmmm. Starting to sound familiar?] Stenomatidae Antaeotricha Stenomatidae Antaeotricha Stenomatidae Rectiostoma

Gelechioidea mini-lab Gelechiidae Smooth scaled head Long upturned labial palpi Proboscis scaled Hindwing with falcate apex (at least pointed) Pattern and shape extremely variable

Sesiioidea (Heteroneura - Ditrysia) 3 families with ca. 1,350 described species Sesiidae, Brachodoidae, Castniidae Woodboring larvae (mostly); some in woody shrubs or woody herbs (e.g., Cucurbita sp.)

Sesiioidea (Heteroneura - Ditrysia) 3 families with ca. 1,350 described species Heteroneurous wings Retinaculo-frenate wing coupling (Sesiidae)

Sesiioidea (Heteroneura - Ditrysia) 3 families with ca. 1,350 described species Heteroneurous wings Retinaculo-frenate wing coupling Ditrysian female reproductive system

Sesiioidea (Heteroneura - Ditrysia) 3 families with ca. 1,350 described species Heteroneurous wings Retinaculo-frenate wing coupling Ditrysian female reproductive system Mouthparts well developed [mini-lab with Tortricoidea]

Tortricoidea (Apoditrysia) 1 family with ca. 9,800 described species Two major subfamilies previously considered families: Olethreutidae Tortricidae Many economically important pests – spruce bud worm, codling moth, light brown applemoth, European grape berry moth

Tortricoidea (Apoditrysia) 1 family with ca. 9,800 described species Heteroneurous wings Frenate wing coupling

Tortricoidea (Apoditrysia) 1 family with ca. 9,800 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system

Tortricoidea (Apoditrysia) 1 family with ca. 9,800 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system Mouthparts well developed

Tortricidae mini-lab Tortricinae Argyrotaenia Tortricinae Choristoneura Labial palpi moderate to short, weakly upturned or porrect Wings with bell-shape silhouette in resting posture (e.g., Archpini) Males of many species with secondary scales (costal fold, densely scaled legs) Most Olethreutinae with costal strigulae on forewing Olethreutinae Eumarozia Tortricinae Clepsis Olethreutinae Grapholita

Sesiidae mini-lab Wasp and bee mimics Usually with clear wings Diurnally active Most easily collected using artificial male pheromones

Zygaenoidea (Apoditrysia) 12 families with ca. 2,700 described species Small to medium sized fuzzy moths (Limacodidae, Megalopygidae, Dalceridae) Bizarre larvae (Limacodidae, Megalopygidae, Dalceridae) Many Zygaenidae are diurnal Epipyropidae are ectoparasites on Homoptera

Zygaenoidea (Apoditrysia) 12 families with ca. 2,700 described species Heteroneurous wings Frenate wing coupling

Zygaenoidea (Apoditrysia) 12 families with ca. 2,700 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system

Zygaenoidea (Apoditrysia) 12 families with ca. 2,700 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system Palpi reduced, proboscis present

Zygaenoidea mini-lab Megalopygidae, Limacodidae, Epipyropidae, and Dalceridae with short, broad wing Zygaenidae with more slenderer body and slenderer wings

Limacodidae larvae – bizarre!

Pterophoroidea (Apoditrysia) 1 family with ca. 1,000 described species Characteristically incised wings Long slender legs Distinctive resting posture

Pterophoridea (Apoditrysia) 1 family with ca. 1,000 described species Heteroneurous wings Frenate wing coupling

Pterophoridea (Apoditrysia) 1 family with ca. 1,000 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system

Pterophoridea (Apoditrysia) 1 family with ca. 1,000 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system Labial palpi variable in shape and vestiture

Pterophoridea (Apoditrysia) 1 family with ca. 1,000 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system Labial palpi variable in shape and vestiture Alucitoidea Alucitidae

Pyraloidea (Obtectomera) 2 families with ca. 17,000 described species

Pyraloidea (Obtectomera) 2 families with ca. 17,000 described species Heteroneurous wings Frenate wing coupling

Pyraloidea (Obtectomera) 2 families with ca. 17,000 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system

Pyraloidea (Obtectomera) 2 families with ca. 1,700 described species Heteroneurous wings Frenate wing coupling Ditrysian female reproductive system Labial palpi variable in shape and vestiture

Pyraloidea mini-lab Superfamily defined by presence of abdominal tympanum; subfamilies frequently defined by various modifications of abdominal tympanum Base of proboscis scaled Most crambids with conspicuous , scaled maxillary palpi