Major patterns in

insect evolution
•What is the relationship between
insects and other arthropods?

•Major steps in insect evolution during

earth’s history
 What is the relationship
between insects and other
arthropods?

- molecular data
- morphology
 morphology says ...
Hexapoda

Myriapoda
Chelicerates
Onycophora Crustacea

Annelid

•insects are most

closely related to
myriapods
but the molecules say....
Crustacea
Myriapoda Hexapoda
Chelicerates

Onycophora
annelid

•insects are most

closely related to
Crustacea
 hexapods and insects
colonized the land
•about 400 mya
•very poor fossil record for the first 60
million years
•about 375 mya, a massive radiation
of insects began
 A major insect
radiation began
in the
Carboniferous
1st radiation

Paleozoic Mesozoic Cenozoic

time
Carboniferous - well-developed
terrestrial ecosystems
a variety of insect orders
who’s on first?

the apterygotes
 the apterygotes

entognath
hexapods

mouthparts
external
 Entognath hexapods

Collembola

Diplura Protura
entognathy
=internal
mouthparts
 the apterygotes

Thysanura

Archeognatha

entognath
hexapods
dicondylic
mandibles

mouthparts
external
 next,
insects evolved wings
the origin of
wings opens
up new ways
of life
In what context did wings evolve?
• soaring from trees
• lobes to protect legs/gills
• thermoregulatory structures
 Stonefly skimming

• nice series of logical steps illustrated in

stoneflies
• maps well onto phylogeny of stoneflies
• http://www.bio.psu.edu/People/Faculty/Marden/
 these are the
oldest kind of
insect wing
-Paleoptera

wings are always completely exposed

wings can’t be folded (flexed) over the back
•the groups ‘above’ the Paleoptera can flex
their wings over their back
• ‘new’ kind of wing = Neoptera
 here are more
Neopteran insects
that originated in the
Paleozoic
 the boundary between the
Paleozoic and Mesozoic is
marked by a great extinction

how well did the different types of

insects do?
 XX XX X X
X X

Insects and especially Paleoptera

were hit hard
X

Most Neopteran orders

survived and many
then radiated in the
Mesozoic
Paleopteran survivors held their own in
the Mesozoic
continued increase in insect diversity
major radiations
in some groups
Hemiptera
plant feeding insects have to eat a
lot of indigestable cellulose

• analogous to a box of candy - and

having to eat the whole box
Hemiptera
piercing/sucking mouthparts
•penetrate plant cells
•penetrate phloem/xylem
•preadaptation for feeding on animal fluids
 Holometabolous

•divergence of larval and adult forms - ecological

niches, specialization of function

•larval stage with internally developing appendages

and crawl inside the candy box

•HUGE increase in potential niches - living in food,

including parasites
 also notice continued
radiation in the Cenozoic

• rise of the angiosperms

• most dominant Holometabola are
associated with flowering plants
• controversial
How did the 3-part life cycle (LPA)
so well known in holometabolous
insects evolve from a
hemimetabolous life cycle (LA)?
 Two hypotheses

1. larva and nymphs are equivalent, and

pupal stage arose to accommodate
increasingly complex transition from larva to
adult (Hinton, 1948)

2. larva is more similar to last embryonic

stage, pronymph, in hemimetabolous insects
(Berlese, 1913)
A A A
A
L P N P
L L N L
L L N L
E E
E E
Hemi Holo Hemi Holo

Hinton Berlese
1999. JW Truman and LM
Riddiford. The origins of
metamorphosis. Nature 401: 447-
452.

2002. Endocrine insights into the

evolution of metamorphosis in
insects. Ann. Rev. Entomol. 47,
467-500.
 Arthropods

•What type of stage hatches from the egg?

•Crustacea - nauplius (non-feeding)

Artemia
 What about insects?

•Apterygota - pronymph,
non-feeding, lasts 3-4 days

•very similar to Crustacea

 What about
hemimetabolous insects?
•Some
Hemimetabola also
have distinct
pronymph

= pharate
1st instar
Odonata, Orthoptera
 Pronymphal stage in
Hemimetabola
•cuticle has distinct ultrastructure - not sclerotized

•lasts several minutes to several hours after hatching

•no wing buds

 Pronymphal stage in
Hemimetabola
• cuticle made at ‘dorsal closure’
katatrepsis and
dorsal closure

Fig. 14.19
• relationship of katatrepsis and dorsal
closure with presence of different cuticle
types in Apterygota and Hemimetabola
How does pronymph compare
to holometabolous larvae?
Look! No nymphal cuticle
How does pronymph compare
to holometabolous larvae?

Similarities

•cuticle with little sclerotization

•larval cuticle secreted at ‘dorsal closure’

Is the holometabolous larva
derived from the
pronymphal stage?

patterns in hormone secretion and

nervous system development also
say yes
 hemi

transition to L

holometaboly

continued
evolution of
holometaboly
 Holometaboly
Hemiptera

Paleoptera Neoptera
Thysanura

Archeognatha wing flexion

entognath wings
hexapods
dicondylic mandibles

mouthparts
external
Major patterns in
insect evolution