The Origin of Complete Metamorphosis: Endopterygota (= Holometabola)

300 250 mya

85%
Crustacea? Myriapoda?
Hexapoda
Collembola
Protura

Diplura
Insecta 15%
Archaeognatha
Dicondylia
Zygentoma
Pterygota
Ephemeroptera
= aquatic immatures Odonata
Neoptera
Plecoptera Polyneoptera**
Paurometabola (orthopteroids)
complete metamorphosis
Endopterygota (Holometabola)
[Eumetabola]
Paraneoptera (hemipteroids)**
(** ‘Exopterygota’)
few malpighian tubules, etc. (weak); furcasternum (better)

A working hypothesis of holometabolan relationships

PANORPIDA

dobsonflies,
alderflies, snakeflies,
lacewings & beetles

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 Pterygota, Neoptera, [Eumetabola], Holometabola: Neuropterodea
Neuropterida (the smaller neuropteroid orders)
Order Megaloptera (270 spp): Corydalidae
dobsonflies and alderflies Order Raphidioptera (210 spp)
snakeflies
adult larva

Ascalaphidae

Order Neuroptera
(6000 spp, 17 fam.)
lacewings, antlions, etc.
Chrysopidae

Nemopteridae
Chrysopidae (Nothochrysa) Ascalaphidae

Myrmeleontidae

Osmylidae
Mantispidae

Sisyridae

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 Pterygota, Neoptera, Eumetabola, Holometabola: Neuropterodea
Order Coleoptera (beetles) 4 suborders, 179 families, ~400,000 spp,
prothorax

elytron

HOLOMETABOLA
in blue:
82-85% of total

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 Some well-known families of beetles, plus the mysterious Strepsiptera

Cantharidae Cerambycidae
(soldier beetles) (long-horned beetles;
(35,000 spp)

Strepsiptera:
Staphylinidae beetle (or fly)?
(rove beetles; 30,000 spp) Curculionidae
(weevils; >60,000 spp)

Scarabaeidae
(scarab beetles;
>30,000 spp)

Homogeneity of gestalt over a vast range of sizes:

Cerambycidae: > 17 cm long

Titanus giganteus
Ptiliidae: barely 0.3 mm long
Nanosella fungi (a staphilinoid)

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 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

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 Despite armored, heavy bodies and life in confined spaces,
Coleoptera are excellent fliers (and they thermoregulate)

Cicindelidae (Carabidae): active, Scarabaeidae: fly immediately &

highly alert predators swiftly to new food sources

Pterygota, Neoptera, Eumetabola, Holometabola: Order Strepsiptera (600 spp)

elytron or
male haltere?
male

female
triungulin
larva

female cephalothorax

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 The other side of the holometabolan tree: Panorpida

flies, fleas, & scorpionflies

PANORPIDA
PANORPIDA
; reduced/lost ovipositor;
CuP+A1 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 Sarcophagidae
(flesh flies) Calliphoridae
(fungus flies) maggots w/mouth hooks
(blow flies)

haltere

Tipulidae
(crane flies)

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 Siphonaptera life cycle,
illustrated

larvae in feces

eggs

adults feeding

pupa
cocoon

Pterygota, Neoptera, Eumetabola, Holometabola: Antliophora –

the haltere of the Diptera

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 Tremendous morphological & biological diversity:

Tipulidae Asilidae
(crane flies) (robber flies)

Streblidae
(bat flies)

Muscidae
(house flies)

Back to basics again: Panorpida

caddisflies, butterflies,
and moths

PANORPIDA
PANORPIDA
; reduced/lost ovipositor;
CuP+A1 fused)

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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.

scales
Arctiidae Pieridae

prolegs

proboscis

Micropterygidae
(Zeugloptera)
Lepidoptera
Agathiphagidae
(Aglossata)

Eriocraniidae
(”Dacnonypha”)
Mnesarchaeidae
(Exoporia)
Aenigmatineidae
Nepticuloidea (Heteroneura)
(”Monotrysia”)
Bombycoidea:
Atlas moth Incurvarioidea Papilionoidea:
(”Monotrysia”) Birdwing butterfly
Palaephatidae
(”Monotrysia”)

Tischeriidae
(”Monotrysia”)

Tineoidea
(Ditrysia)
Yponomeutoidea
(Ditrysia)
Gelechioidea
(Ditrysia)

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 Back to basics again: Hymenopterodea

sawflies, wood wasps,

wasps, ants, & bees

PANORPIDA
PANORPIDA
; reduced/lost ovipositor;
CuP+A1 fused)

Pterygota, Neoptera, [Eumetabola], Holometabola: Hymenoptera

(sawflies, bees, wasps & ants) – 180,000 spp in 91 fam.

s.o. Symphyta: s.o. Apocrita:

(incl. Parasitica & Aculeata)
Formicidae
Apidae
Orussidae

Vespidae

Ichneumonidae

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 s.o. Symphyta
Sawflies Wood wasps

Cicadidae (Hemiptera)
Tettigoniidae (Orthoptera)

Ichneumonidae
(Hymenoptera) Apidae (Hymenoptera)

Thermobia
(Zygentoma)
Muscidae (Diptera)
(functional
Evolution of the
“ovipositor”)
hymenopteran stinger
in the s.o. Apocrita

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 Dolichovespula maculata
Key adaptations:
The hymenopteran body is hard to crush;

…but the key to the success

of the Hymenoptera is eusociality:
Honeybee (Apis mellifera)

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