Unlike
the trilobite that has left
a prodigious fossil record,
the preservation of insects in sedimentary matrix is relatively
rare, and essentially limited to the Laggerstat sites. The reason
for the scarcity of insect fossil is the poor preservation potential
of the insect's exoskeleton. Like other Arthropods, insects
have an external skeleton called an exoskeleton. Unlike the
thick and calcified trilobite exoskeleton, the insect exoskeleton
is made of a thin, plastic-like material called chitin, along
with a tough protein. This thin, waterproof covering simple
does not preserve well in most oxygenated environments, making
insect fossils sparse despite the tremendous number that could
have been preserved. The exception is in fossil
resinite (amber, by street name), where it is possible for
even the minutest details to be preserved. Despite their huge
strength to weight ratio, insects were often too small to escape
the sticky resin exuded by trees, and which later became a fossil
itself, with physical properties akin to modern polymerized
plastics. Countering low probability of fossilization is the
prodigious fecundity of insects. While lifetime insect fecundity
varies enormously across taxa, from less than ten to several
millions of eggs, most insects lay between a couple of hundred
to a couple of thousand eggs (Hinton
1981).
Insect
evolution is a powerful illustration of decent with modification.
The earliest known insects are tiny wingless forms from the
early and middle Devonian.
Insect flight developed with suddenness resembling the Cambrian
explosion during the middle Carboniferous, apparently the result
of the significant survival advantage that was accrued. By the
end of the Carboniferous, the subphylum insecta had evolved
into a large number of distinct orders. During the Permian,
new insects forms appeared. Blattoid and Orthopteroid orders
attained their greatest diversity, and new groups like the Psocoptera,
Homoptera, Hemiptera, Mecoptera and Coleoptera
became ubiquitous and diverse. The Permian extinction wiped
out nine orders of insects, and more orders disappeared in the
Triassic or the early Jurassic. However, surviving orders such
as Neuroptera, Mecoptera, and Diptera, and Coleoptera underwent
further adaptive radiation establishing many families extant
in modern times. So exquisite is insect design that most groups
were well formed by the Cretaceous and remain largely unchanged
in appearance during modern times.
Insect
evolution has led to prodigious diversity of this animal group
comapred with other members of Domain
Eukarya. For example, there are believed to be three times
as many Dipteran
species (true fllies) as there are vertebrate species, and ten
times as many Coleopteran
species (beetles) as there are vertebrate species.
Taxonomic
research on fossil insects has always been relegated to a subordinate role when
compared to that of living species. There are large numbers of undetermined fossil
insects in many collections throughout the world awaiting descriptions, but only
a small fraction of systematic research has ever been devoted to these fossils.
Regarding
insect origins, Gaunt and Miles
(2002) propose based molecular clock Analyses encompassing five
insect orders (Blattaria, Orthoptera, Hemiptera, Diptera, and
Lepidoptera) that the class insecta common ancestor may have
been Anostraca (fairy shrimps) at around the Ordovician-Silurian
boundary around 434 to 421 million years ago. This time coincides
with the earliest plant megafossil, and is consistent with the
long held view that terrestrial transition of the aquatic arthropod
ancestor to the insects is associated with the appearance with
early
vascular plants.
Subphylum
Hexapoda - Class
Insecta and Class Entognatha
| H
E
X
O
P
O
D
A
|
Class |
Subclass |
Infraclass |
Superorder |
Order
(Links
are to Fossils by Order) |
Common
Names
within order |
|
Approximate
Extant species described |
| Entognatha
(2) |
Collembola |
n/a |
|
Collembola |
Springtails |
Devonian |
2,000 |
| |
n/a |
|
Diplura |
Two-pronged
bristletails |
Carboniferous |
800 |
| |
n/a |
|
Protura |
Coneheads |
Devonian |
rare/100 |
| Insecta |
Apterygotes
- never had wings (1) |
n/a |
|
Archaeognatha |
Jumping
bristletails |
Devonian |
350 |
| n/a |
|
Monura |
Enigmatic
taxon |
Carboniferous |
Extinct |
| n/a |
|
Thysanura |
Silverfish |
Lower
Devonian |
700 |
| |
|
|
|
|
|
|
| |
Ephemeroptera |
Mayfly |
Devonian |
2,100 |
| |
Odonata |
Dragonflies;
Damselflies |
Devonian |
>5,500 |
| Pterygota
(Have or had wings) |
Neoptera |
Exopterygota |
Blattaria |
Cockroachs |
Mississippian |
3,700 |
| Mantodea |
Mantid |
Pennsylvannian |
>1,800 |
| Mantophasmatodea |
Gladiator |
- |
- |
| Isoptera |
Termite |
Upper
Cretaceous |
2,000 |
| Plecoptera |
Stonefly |
Permian |
1,600 |
| Orthoptera |
grasshoppers,
crickets and locusts |
Mississippian |
20,000 |
| Embioptera
|
webspinners |
? |
300 |
| Dermaptera |
earwigs |
Jurrasic |
2,000 |
| Phthiraptera |
lice |
? |
3000 |
| Notoptera
|
ice-crawlers
& gladiators |
? |
<30 |
| Embioptera |
webspinners |
? |
170 |
| Zoraptera |
angel
insects |
? |
1
genus |
| Phasmatodea |
Walking
stick; walking leaf |
Lower
Triassic |
2,500 |
| Thysanoptera |
thrips |
Permian |
5000 |
| Psocoptera |
booklice,
barklicee |
Permian |
6,000 |
| Hemiptera |
true
bugs: Cicada; aphid; plant hopper; leaf hopper; spittlebugs;
scale insects; mealy bugs |
Upper
Pennsylvannian |
82,000 |
| Endopterygota |
Hymenoptera |
ants;
bees; wasps; sawflies |
Upper
Triassic |
130,000 |
| Homoptera
(suborder combined with Hemiptera) |
cicadas;
aphids; plant hoppers; leaf hoppers; spittlebugs; scale
insects; mealy bugs |
Permian |
33,000 |
| Coleoptera |
Beetles |
Lower
Permian |
350,000 |
| Neuroptera |
Lacewings;
antlions |
Lower
Permian |
4,700 |
| Strepsiptera |
twisted-winged
parasites |
? |
? |
| Raphidioptera
|
snakeflies |
Jurassic |
150 |
| Mecoptera |
Scorpianflies |
Pennsylvannian |
550 |
| Siphonoptera |
fleas |
Miocene
(?) |
1,750 |
| Diptera |
mosquitos,
gnats, midges, etc. |
Middle
Triassic |
240,000 |
| Trichoptera |
caddisflies |
Lower
Triassic |
7,000 |
| Lepidoptera |
Butterfly;
moth |
Jurassic |
180,000 |
| (1)
The taxon Apterygota is often applied for a subclass of
insects lacking wings in both extant forms as well across
their evolutionary history. The earliest occurence in
the fossil record is the Devonian period some 417-354
million years ago. The nymphs go through little or no
metamorphosis, and thus closely resemble the adults. Apterygotans
generally have a thin exoskeleton, giving them a pseudo-translucent
characteristic.
(2)
The Entognatha is a class of arthropods, which, together
with insects, makes up the hexapods. They exhibit little
or no metamorphosis. The class's three orders: Collembola,
Diplura and Protura are currently believed to have independently
evolved six legs, and to have independently evolved within
each order, making the Entognatha a polyphyletic group.
(3)
The Palaeoptera comprises a primitive grouping of winged
insects (mostly extinct forms) that lack or lacked the
ability to fold the wings back over the abdomen as members
of Neoptera do. The relationship of the two extant Paleoptera
groups (Ephemeroptera and Odonata) to the Neoptera is
unresolved; some believe Paleoptera to be paraphyletic
such that the grouping should be abandoned. |
|