Exoskeleton vs.

Endoskeleton: biomechanical trade-offs

Ratio:
surface area (square)
. . volume (cube)

strength of skeleton
Hercules beetle
(Dynastes hercules)
Shrew (Sorex sp.)

size of organism
a giant spider

Exoskeleton vs. Endoskeleton

The exception proves the rule, in aquatic arthropods.

…but this is not

possible)

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 Other advantages of the exoskeleton
The necessity of molting allows
complete metamorphosis

The cuticle includes

a lipid monolayer

The cuticle is impact resistant

What about six legs??

“Hexapoda” may not be monophyletic

Crustacea? Myriapoda?
Hexapoda (six legs)
Collembola
Protura

Diplura
Insecta
Archaeognatha
Dicondylia
Zygentoma
Pterygota
Ephemeroptera
Odonata
Neoptera
Plecoptera Polyneoptera**
Paurometabola (orthopteroids)
Endopterygota (Holometabola)
Eumetabola
Paraneoptera (hemipteroids)**
(** ‘Exopterygota’)

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 Three classes of Entognathous hexapods:
Collembola, Protura, Diplura

Class Collembola (springtails):

Class Diplura:

oral cone
(collophore)

Class Protura:
12

Entognatha: Class Collembola (springtails, snow fleas)

6000 species
12-20 families
4 subclasses
(Lower Devonian)

Not a true insect:

• 6-segmented
abdomen

3
 Entognatha: Class Protura (“proturans”)

730 species
3-8 families
2 subclasses
(Devonian)

Not an insect:
• no antennae
• anamorphosis

Entognatha: Class Diplura

(“diplurans”)

800 species
5-9 families
Not an insect: 2 subclasses
• antennal flagellum w/ (Carboniferous)
intrinsic muscles
• glandular cerci

4
 The three major evolutionary innovations of the Class Insecta:
wings, wing-folding, & complete metamorphosis (≈ 27 orders, total)

Crustacea? Myriapoda?
Hexapoda
Collembola
Protura

Diplura
Insecta (Ectognatha)
Archaeognatha
Dicondylia
Zygentoma
1. Pterygota
Ephemeroptera
= aquatic immatures Odonata
2. Neoptera
Plecoptera Polyneoptera**
Paurometabola (orthopteroids)
complete metamorphosis
3. Endopterygota (Holometabola)
Eumetabola
Paraneoptera (hemipteroids)**
(** ‘Exopterygota’)

3 ocelli
order Archaeognatha (= Microcoryphia, Machiloidea)
compound eye
antenna
cercus

median caudal filament (MCF)

labial abdominal styli
palp (seg. 2-9)
(3-seg)
maxillary palp style on
coxa ovipositor
(7-segmented)
exsertile vescicle
incisor region
order Zygentoma
(Thysanura) mandibles
maxillary palp
molar
region
(mono- vs dicondylic)
condyle

incisor region molar region

condyle

condyle

style (8 & 9)
style ovipositor
cercus cercus

median
‘subclass Apterygota’
caudal
filament median
caudal
filament

5
 Apterygota:
Archaeognatha (bristletails)
350 species, 2 families; Devonian (?)

leg-like maxillary & labial palps

Monura?

median caudal filament

5 mm
Permian Dasyleptidae (Elmo fauna, Kansas)

Families: Meinertellidae & Machilidae

Apterygota: order Zygentoma (silverfish & firebrats)

370 species, 3-5 families 2 condyles

Thermobia sp.

Lepisma sp.

Tricholepidion gertschi
lives w/ w.NA. carpenter ants)

6
 Apterygota are probably paraphyletic,
not monophyletic
(‘Apterygota’ (blue bar) is probably not a clade)

outgroup (monocondylic)

?
Monura (extinct; condyly uncertain)
Archaeognatha (=Microcoryphia)
dicondylic
mandible
Zygentoma (=Thysanura)
Dicondylia
Pterygota (other insects)

• monophyletic: an ancestral species and all its descendants (a clade).

• paraphyletic: a monophyletic group from which one or more
subsidiary clades (monophyletic groups) is excluded.

The origin of wings: Paleoptera

Crustacea? Myriapoda?
Hexapoda
Collembola
Protura

Diplura
Insecta
Archaeognatha
Dicondylia
Zygentoma
Pterygota
Ephemeroptera
= aquatic immatures Odonata
Neoptera
Plecoptera Polyneoptera**
Paurometabola (orthopteroids)
complete metamorphosis
Endopterygota (Holometabola)
Eumetabola
Paraneoptera (hemipteroids)**
(** ‘Exopterygota’)

7
 The origin of wings: Paleoptera
Ancestral groups all have aquatic immatures:
Holometabola
Paleoptera Neoptera
Ephemeroptera Odonata Plecoptera Megaloptera

Pterygota, Paleoptera:
Ephemeroptera (mayflies)
> 3000 species, 25 families; suborders & superfamilies uncertain
Adult
adult Naiad (immature)
naiad (nymph)

Ancestral features:
• 11-segmented abdomen
• cerci & median caudal
filament
• 3 ocelli
• post-adult molt
• paired genitalia (!)
(as in Archaeognatha & Zygentoma)

8
 Mayflies – aquatic immatures

Nymph/naiad:
• subaquatic
• long-lived (vs. adult)
• tracheal gills
• morphologically
diverse

Permian mayflies: Protereisma sp. (Protereismatidae)

long antennae 5-segmented

Naiad (immature) tarsus
(after Kukalova) nearly full
wing venation
subequal
wings

Adult
(after Carpenter)
9 pairs
of gills

10th

MCF

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 Pterygota, Paleoptera:
Odonata (dragonflies & damselflies)
Two (or three) suborders, 27 families, 600 genera, 6500 extant species

‘s.o.’ Anisoptera ‘s.o.’ Zygoptera

(dragonflies) (damselflies)

caudal
gills

Thoracic tilt

s.o. Anisoptera
(dragonflies)
s.o. Zygoptera
(damselflies)

10
 Tracheal gills and nymphal “mask”

Anisoptera naiad (rectal gills)

“mask”

Zygoptera (caudal gills)

“mask”

Damselflies

11
 Dragonflies Libelluloidea

Aeshnoidea

Dragonfly progenitors: order Protodonata

(Carboniferous, approx 350 mya)
28 inches (70 cm)

Meganeuridae: Meganeura

Meganeuridae: Megatypus

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 Phylogenetic hypotheses for Paleoptera

I. Paleoptera are monophyletic (Hennig)

Outgroup
Neoptera (all other winged insects)
Odonata
Paleoptera
Ephemeroptera

II. Paleoptera are paraphyletic

(Kristensen) (Boudreaux)
Outgroup Outgroup
Ephemeroptera Odonata
Paleoptera Paleoptera
Odonata Ephemeroptera
Neoptera (all other winged insects) Neoptera (all others)

The Origin of Wing-Folding: Neoptera

Crustacea? Myriapoda?
Hexapoda
Collembola
Protura

Diplura
Insecta
Archaeognatha
Dicondylia
Zygentoma
Pterygota
Ephemeroptera
= aquatic immatures Odonata
Neoptera
Plecoptera Polyneoptera**
Paurometabola (orthopteroids)
complete metamorphosis
Endopterygota (Holometabola)
Eumetabola
Paraneoptera (hemipteroids)**
(** ‘Exopterygota’)

13
The morphological basis of wing-folding (neoptery): Axillary sclerites

Paleoptera
PALEOPTERA(no(no
wing-folding)
wing-folding)
C+
Sc-
R+
Rs-
MA+
MP-
CuA+
CuP-

2A+ 1A+ Odonata

Neoptera
NEOPTERA(wing-folding)
(wing folding)
3A+

C+
Sc-
R+
Rs-
MA+
MP-
CuA+
posterior notal process 1A+
2A+ Plecoptera
3A+

Pterygota, Neoptera, Polyneoptera: order Plecoptera (stoneflies)

3500 species, 2 ‘suborders’, 16 families
adult:
adult:

naiad:
naiad
(nymph)
(immature)

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 Pterygota, Neoptera, Polyneoptera:
Order Grylloblattodea (ice or rock crawlers)
> 33 species, 5 genera, 1 family (Grylloblattidae)

Discovered in 1913 ovipositor

(Walker 1914) as a cercus
“living fossil”)

Pterygota, Neoptera, Polyneoptera:

Order Mantophasmatodea (gladiators, heelwalkers)
~24 living species, 3 families, ≥ 8 genera

(first recognized
in 2002!)

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 Pterygota, Neoptera, Polyneoptera:
order Dictyoptera, s.o. Blattodea (cockroaches)
4500 species, 6–9 families

Blattella germanica

Cockroaches, past & present:

Periplaneta sp.
R
Sc
C
M
“Blattoptera:”
Archimylacridae
(Carboniferous)
Sc
R1
CuA
C Sc CuP R
Rs

M
M
CuA 1A CuA
CuP CuP

anal fan 300 my-old fossil, 4” long

Carboniferous (central Asia)

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Pterygota, Neoptera, Polyneoptera: order Dictyoptera, s.o. Isoptera (termites)
3100 species, 7 families (all eusocial)

“isopterous” wings

workers

nests (colonies)

Mastotermes darwiniensis (Mastotermitidae)

Termites:
“Social cockroaches”
imago

eggs in ootheca

anal lobe

Indotermes soldier
(Termitidae)

Amitermes hastatus
Synapomorphies: (primitiveTermitidae)
1. Eusociality
2. True castes

worker tertiary queen 2ndary queen primary queen

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 Pterygota, Neoptera, Polyneoptera:
order Dictyoptera, s.o. Mantodea (mantises)
2400 species, 15 families

ootheca

Pterygota, Neoptera, Polyneoptera:

order Phasmatodea (stick & leaf insects) order Dermaptera (earwigs)

3000 species, 2 suborders, several (?) families 2200 species, 3 ‘suborders,’ 11 families
s.o. Hemimerina

s.o. Forficulina

order Embioptera (webspinners) s.o. Arixenina

360 species, 2 suborders, 8 families
Timema sp.

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 Pterygota, Neoptera, Polyneoptera:
Order Orthoptera (grasshoppers, crickets & katydids)
22,000 species, 2 suborders, many families
suborder Ensifera (11,000 spp):
suborder Caelifera (11,000 spp): crickets, katydids, long-horned grasshoppers
Acrididae (short-horned grasshopper)

ear

ear

Specialized Neoptera without complete metamorphosis:

the Paraneoptera

Crustacea? Myriapoda?
Hexapoda
Collembola
Protura

Diplura
Insecta
Archaeognatha
Dicondylia
Zygentoma
Pterygota
Ephemeroptera
= aquatic immatures Odonata
Neoptera
Plecoptera Polyneoptera**
Paurometabola (orthopteroids)
complete metamorphosis
Endopterygota (Holometabola)
Eumetabola
Paraneoptera (hemipteroids)**
(** ‘Exopterygota’)

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 Pterygota, Neoptera, [Eumetabola], Paraneoptera:
the smaller “hemipteroid” orders
(orthopteroid) order order Thysanoptera: thrips
Zoraptera ‘Psocoptera:’ book lice (5500 spp) (6000 spp)
(39 spp)

order
Psocodea:
lice
(11,000 spp)

‘Phthiraptera:’
‘s.o.’ Mallophaga, ‘s.o.’ Anoplura,
biting lice (4450 spp) sucking lice (550 spp)

Pterygota, Neoptera, [Eumetabola], Paraneoptera:

more thrips (Thysanoptera)

soldier caste, Carcinothrips leai

(Australia)

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 Pterygota, Neoptera, Paraneoptera: Order Hemiptera (bugs)
3 (or 4) suborders, >133 families, >2000 genera, 100,000 species
(size: 1mm - 110mm!)

s.o. Auchenorrhyncha (40,000 spp): cicadas and hoppers

Membracidae: Cicadellidae: Cicadidae: cicadas
tree hoppers leaf hoppers

s.o. Sternorrhyncha (25,000 spp):

aphids, scales, whiteflies, other plant lice

Fulgoroidea: plant hoppers

s.o. (?) Coleorrhyncha s.o. Heteroptera (30,000 spp): “true bugs”

(34 spp) Peloridiidae –
Gondwanan distrib.
Lygaeidae (seed bugs) Coreidae (squash bugs) Pentatomidae (stink bugs)

Miridae (plant bugs)

(heteropterous wing)

Nepidae (water bugs)

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