The Origin of Complete Metamorphosis: Endopterygota (= Holometabola) Hexapoda 300 250 mya Crustacea? Myriapoda? Collembola Protura 85% Insecta = aquatic immatures (** Exopterygota ) Dicondylia Pterygota Neoptera complete metamorphosis [Eumetabola] Diplura Archaeognatha Zygentoma Ephemeroptera Odonata Plecoptera Polyneoptera** Paurometabola (orthopteroids) Endopterygota (Holometabola) Paraneoptera (hemipteroids)** few malpighian tubules, etc. (weak); furcasternum (better) 15% A working hypothesis of holometabolan relationships dobsonflies, alderflies, snakeflies, lacewings & beetles PANORPIDA 1
Pterygota, Neoptera, [Eumetabola], Holometabola: Neuropterodea Neuropterida (the smaller neuropteroid orders) Order Megaloptera (270 spp): Corydalidae dobsonflies and alderflies adult larva Order Raphidioptera (210 spp) snakeflies Order Neuroptera (6000 spp, 17 fam.) lacewings, antlions, etc. Chrysopidae Ascalaphidae Nemopteridae Chrysopidae (Nothochrysa) Ascalaphidae Myrmeleontidae Mantispidae Osmylidae Sisyridae 2
Pterygota, Neoptera, Eumetabola, Holometabola: Neuropterodea Order Coleoptera (beetles) 4 suborders, 179 families, ~400,000 spp, prothorax elytron HOLOMETABOLA in blue: 82-85% of total 3
Some well-known families of beetles, plus the mysterious Strepsiptera Cantharidae (soldier beetles) Cerambycidae (long-horned beetles; (35,000 spp) Staphylinidae (rove beetles; 30,000 spp) Curculionidae (weevils; >60,000 spp) Strepsiptera: beetle (or fly)? Scarabaeidae (scarab beetles; >30,000 spp) Homogeneity of gestalt over a vast range of sizes: Ptiliidae: barely 0.3 mm long Nanosella fungi (a staphilinoid) Cerambycidae: > 17 cm long Titanus giganteus 4
Big or little, beetles pretty much look like beetles Histeridae: Bacanius punctiformis Cerambycidae: Titanus giganteus (6.5 inches long; rain forests of South America) But not their larvae! Dytiscidae Scarabaeidae Chrysomelidae Elateridae Coccinellidae Curculionidae 5
Despite armored, heavy bodies and life in confined spaces, Coleoptera are excellent fliers (and they thermoregulate) Cicindelidae (Carabidae): active, highly alert predators Scarabaeidae: fly immediately & swiftly to new food sources Pterygota, Neoptera, Eumetabola, Holometabola: Order Strepsiptera (600 spp) male elytron or haltere? male female triungulin larva female cephalothorax 6
The other side of the holometabolan tree: Panorpida flies, fleas, & scorpionflies PANORPIDA ; reduced/lost ovipositor; CuP+A 1 fused) Pterygota, Neoptera, [Eumetabola], Holometabola: Antliophora Order Mecoptera (scorpionflies) 600 spp Order Siphonaptera (fleas) 2575 spp Bittacidae rostrum Panorpidae Order Diptera (true flies) 154,000 spp in 151 fam. Mycetophilidae (fungus flies) Sarcophagidae (flesh flies) Calliphoridae (blow flies) maggots w/mouth hooks haltere Tipulidae (crane flies) 7
Siphonaptera life cycle, illustrated larvae in feces eggs adults feeding pupa cocoon Pterygota, Neoptera, Eumetabola, Holometabola: Antliophora the haltere of the Diptera 8
Tremendous morphological & biological diversity: Tipulidae (crane flies) Asilidae (robber flies) Streblidae (bat flies) Muscidae (house flies) Back to basics again: Panorpida caddisflies, butterflies, and moths PANORPIDA ; reduced/lost ovipositor; CuP+A 1 fused) 9
Pterygota, Neoptera, [Eumetabola], Holometabola: Amphiesmenoptera (female is the heterogametic sex, i.e., XZ) Order Trichoptera (caddisflies) 15,000 spp hairs (setae) Larval cases Order Lepidoptera (moths and butterflies) 180,000 spp in 128 fam. Arctiidae scales Pieridae prolegs proboscis Micropterygidae (Zeugloptera) Agathiphagidae (Aglossata) Lepidoptera Eriocraniidae ( Dacnonypha ) Bombycoidea: Atlas moth Mnesarchaeidae (Exoporia) Nepticuloidea ( Monotrysia ) Incurvarioidea ( Monotrysia ) Palaephatidae ( Monotrysia ) Tischeriidae ( Monotrysia ) Aenigmatineidae (Heteroneura) Papilionoidea: Birdwing butterfly Tineoidea (Ditrysia) Yponomeutoidea (Ditrysia) Gelechioidea (Ditrysia) 10
Back to basics again: Hymenopterodea sawflies, wood wasps, wasps, ants, & bees PANORPIDA ; reduced/lost ovipositor; CuP+A 1 fused) Pterygota, Neoptera, [Eumetabola], Holometabola: Hymenoptera (sawflies, bees, wasps & ants) 180,000 spp in 91 fam. s.o. Symphyta: Orussidae Formicidae s.o. Apocrita: (incl. Parasitica & Aculeata) Apidae Vespidae Ichneumonidae 11
Sawflies s.o. Symphyta Wood wasps Cicadidae (Hemiptera) Tettigoniidae (Orthoptera) Ichneumonidae (Hymenoptera) Apidae (Hymenoptera) Thermobia (Zygentoma) Muscidae (Diptera) (functional ovipositor ) Evolution of the hymenopteran stinger in the s.o. Apocrita 12
Key adaptations: The hymenopteran body is hard to crush; Dolichovespula maculata but the key to the success of the Hymenoptera is eusociality: Honeybee (Apis mellifera) 13