FOSSIL INVERTEBRATES/Insects 295
and worldwide. As trilobites dominated the faunas of
the Cambrian and are rich in morphological features,
they are the obvious choice for characterizing local
and regional successions. The Early Cambrian in
Morocco, Siberia, and China is divided into about
10 trilobite biozones in each area; likewise, the
Middle Cambrian of Australia and China is divided
into some 8–10 biozones. In the Late Cambrian of
Australia, 17 trilobite zones are recognized, and
about 15 in Kazakhstan. In the olenid facies of
Scandinavia, there are eight zones divided into about
30 subzones. Correlation between the successions
around different continents is difficult, especially in
the earlier Cambrian, but becomes more secure in the
later half of the Cambrian. The occasional taxa that
crossed the barriers between continents (e.g., Irving-
ella and some Agnostida, such as Glyptagnostus) are
important because they provide key correlative ties
(Figure 18). In the Ordovician and Silurian, grapto-
lites are used widely for intercontinental correlation,
but the pelagic trilobite, Carolinites (Figure 14A), is of
equal value in the earlier Ordovician. Other trilobites
retain their value in local successions, especially those
with few graptolites, e.g., the Early Ordovician of the
Laurentian platform and the Middle Ordovician of
Baltica. Elucidation of the complex patterns of pits
in the fringes of trinucleid trilobites has enabled
detailed correlations locally, e.g., in the Middle and
Late Ordovician of Britain. Trilobites also have bio-
stratigraphical value in certain facies of the Devonian
and Carboniferous.
Insects
E A Jarzembowski, University of Reading, Reading,
UK and Maidstone Museum and Bentlif Art Gallery,
Maidstone, UK
ß 2005, Elsevier Ltd. All Rights Reserved.
Introduction
Insects, which belong to the taxonomic group
Hexapoda (‘six legs’; Table 1), are the most successful
organisms on Earth, if biodiversity is measured as
a count of the number of species. More than
1.4 million species of insects have been described in
the past 250 years, comprising 65% of all known
species of life on Earth (Figure 1A). Insects are also
the most successful group in the fossil record, if
palaeodiversity is measured as a count of the number
of families documented, as many palaeontologists
do (Figure 2).
See Also
Biozones. Evolution. Fossil Invertebrates: Arthropods.
Palaeoecology. Palaeozoic: Cambrian; Ordovician; Si-
lurian; Devonian; Carboniferous; Permian. Trace Fossils.
Further Reading
Clarkson ENK (1979) The visual systems of trilobites.
Palaeontology 22: 1–22.
Fortey RA (2000) Trilobite! Eyewitness to Evolution.
London: HarperCollins.
Fortey RA and Owens RM (1999) Feeding habits in trilo-
bites. Palaeontology 42: 429–465.
Harrington HJ, Henningsmoen G, Howell BF, et al. (1959)
Trilobita. In: Moore RC (ed.) Treatise on Invertebrate
Paleontology, Part O, Arthropoda 1. Lawrence, KS: Geo-
logical Society of America and University of Kansas
Press.
Jell PA and Adrain JM (2003) Available generic names
for frilobites. Memoirs of the Queensland Museum 48:
331–553.
Levi-Setti R (1993) Trilobites, 2nd edn. Chicago and
London: University of Chicago Press.
Šnajdr M (1990) Bohemian Trilobites. Prague: Czech Geo-
logical Survey.
Whittington HB (1992) Fossils Illustrated 2. Trilobites.
Woodbridge: Boydell Press.
Whittington HB, Chatterton BDE, Speyer SE, et al. (1997)
Treatise on Invertebrate Paleontology, Part O, Arthro-
poda 1, Trilobita, revised, vol. 1. Boulder, CO and Law-
rence, KS: Geological Society of America, Inc., and
University of Kansas Press.
How many insects are there? Nobody knows
for sure, although all estimates suggest that millions
of insect species remain to be described. The great
majority of these live in exotic places. It seems un-
likely that description will keep pace with global
habitat loss and extinction brought about by human
Table 1 Insects – systematic position
Taxonomic division Group/number
Kingdom Animalia
Phylum Arthropoda
Superclass Hexapoda (insects in the
broad sense)
Order About 43
Family Over 1500
Genus ?
Species 3–20 million
296 FOSSIL INVERTEBRATES/Insects
Figure 1 Global biodiversity, showing percentage breakdown
for insects and other organisms. (A) Percentages of the app-
roximate number of known species worldwide (1 454 000).
(B) Percentages of the estimated total number of species
worldwide, which includes those thought to be undiscovered
(65 654 000).
development. Simple extrapolation from the fossil
record suggests that the total number of insect species
is probably less than 20 million, although some biolo-
gists prefer a higher figure. Everyone is agreed, never-
theless, that insects represent over 50% of all known
species and that they belong to an exclusive group of
hyperdiverse organisms.
Origins
Where do insects come from? Insects are undoub-
tedly a class of arthropods, or ‘joint-legged’ animals
(Table 1). Arthropods (see Fossil Invertebrates:
Arthropods) also include trilobites, crustaceans
(prawns, etc.), chelicerates (spiders, etc.), and myria-
pods (millipedes and centipedes). Insects show closest
relationships to crustaceans (e.g., in the structure of
their compound eyes) and myriapods (e.g., in their
tubular or tracheate respiratory system) (Figure 3).
The exact relationships are currently a subject of
debate.
Classification
Insects are divided into two main groups – winged
and wingless hexapods (Figure 4). The wingless
(apterygotan) insects are a mixed group and only
some (silverfish) are thought to share a common
ancestor with winged (pterygotan) insects. The
pterygotes are divided into two main groups
(Figure 5) – those that can fold their wings over the
body (Neoptera) and those that cannot (Paleoptera).
The neopterans, in turn, can be divided into two
groups – those that undergo complete metamorphosis
(Holometabola) and those that undergo incomp-
lete metamorphosis (cockroach and grasshopper
orders and bug orders, or Polyneoptera and Para-
neoptera, respectively); the Polyneoptera and
Paraneoptera are also known as exopterygotes, be-
cause the wings develop on the outside in the young
stages (Figure 6). In contrast, the wings develop
inside holometabolous insects, or endopterygotes.
For holometabolans, the chrysalis, or pupa, is the
‘resting’ stage between the caterpillar, maggot, or
grub stage and the flying adult stage. The holometa-
bolans are the most diverse insects and apterygotes
are the least diverse (Figure 7). Some pterygotes have,
however, lost their wings (e.g., fleas). Very high diver-
sities (100 000 or more species) are reached in only
four (holometabolous) orders: Coleoptera (beetles),
Lepidoptera (moths and butterflies), Hymenoptera
(wasps, ants, and bees), and Diptera (true flies).
Geological History
The origin of insects is a mystery, the Cambrian
Burgess Shale arthropods being too early to cast any
light on the subject. The oldest definite hexapod is
Rhyniella praecursor from the Early Devonian Rhynie
Chert. Rhyniella praecursor is a springtail belonging
to the living apterygote order Collembola. The earli-
est true insect is currently considered to be Rhyniog-
natha hirsti, also from the Rhynie Chert. In the latest
Lower Carboniferous and Upper Carboniferous there
is evidence of the radiation of the pterygotes, includ-
ing paleopterans and polyneopterans. These insects
were the world’s first flying animals, long before ver-
tebrates took to the air. In the succeeding Permian, the
paraneopterans and holometabolans became estab-
lished. Insects are essentially terrestrial organisms,
but the first definite freshwater forms appeared in
the Permian. After a setback in the Early Triassic
extinction (Figure 8), insects regained their ordinal
strength by the Tertiary, establishing some new innov-
ations on the way, e.g., evolving parasitic and para-
sitoid forms as well as insect societies (Figure 9). The
extinction at the start of the Mesozoic seems to have
 FOSSIL INVERTEBRATES/Insects 297

Figure 2 Palaeodiversity of organisms. Key to numbers: 1, bacteria and blue-green algae; 2, fungi; 3, other algae; 4, single-cell
organisms; 5, sponges; 6, corals, etc.; 7, chitons, etc.; 8, snails; 9, nautiluses; 10, ceratites; 11, ammonites; 12, belemnites; 13, bivalves
and tusk shells; 14, uncertain molluscs; 15, segmented worms; 16, trilobites; 17, spiders, etc.; 18, crustaceans (excluding seed
shrimps); 19, seed shrimps; 20, millipedes, etc.; 21, insects; 22, lamp shells; 23, phoronids; 24, moss animals; 25, sea urchins, etc.;
26, primitive chordates; 27, graptolites; 28, problematica; 29, miscellaneous; 30, conodonts; 31, lampreys, etc.; 32, cyclostomes; 33,
primitive fish; 34, sharks, etc.; 35, primitive bony fish; 36, advanced bony fish; 37, more bony fish; 38, amphibians; 39, reptiles; 40, birds;
41, mammals; 42, mosses, etc.; 43, ferns, etc.; 44, seed plants (excluding 45, flowering plants).

Figure 3 Relationship of insects with other arthropods.

Figure 5 The two main groups of pterygotes are the paleopter-

Figure 4 Major insect groups; apterygotes include springtails
and silverfish.
ans and the neopterans, which are further divided into subgroups.
Paleopterans include dragonflies and mayflies. Cockroaches and
grasshoppers, along with stoneflies, stick insects, crickets,
locusts, earwigs, termites, and praying mantises comprise the
polyneopteran orders. Paraneopteran orders include bugs and
lice. Holometabolous orders include beetles, lacewings, wasps,
ants, bees, caddisflies, moths, butterflies, flies, and fleas.
298 FOSSIL INVERTEBRATES/Insects
Figure 6 Life cycles of insects. (A) Endopterygotes undergo a
complete metamorphosis. (B) Exopterygotes undergo an incom-
plete metamorphosis.
Figure 7 Approximate biodiversity of major insect groupings,
showing percentages of primitively wingless insects (aptery-
gotes), primitively winged insects with incomplete metamorphosis
(paleopterans and exopterygote neopterans), and primitively
winged insects with complete metamorphosis (holometabolous
insects, or endopterygote neopterans).
been the biggest in insect history, although losses
were not really catastrophic. Indeed, the successful
order Diptera (true flies), which were opportunists
(‘specials’), arose in the Triassic (Figure 8). The
extinction showed, however, that even hyperdiverse
Figure 8 Insect orders during the (A) Triassic extinction; (B)
Phanerozoic composition. Opportunists are categorized as
‘specials’.
organisms are affected by global environmental
change. The principal groups (orders) of insects in
the fossil record are outlined in Table 2.
Collecting and Documentation
Fossil insects are more common than is generally
supposed, especially their disarticulated remains.
They occur in a variety of sedimentary environments,
including marine and non-marine deposits, in both
organic and fine-grained clastic rocks. Insects often
occur in early diagenetic concretions, including
ferruginous and phosphatic ones, and in calcareous
mudstones. Some of the best preserved insects
occur in amber from the Lower Cretaceous onwards.
More unusual modes of preservation include pyrit-
ized or silicified insects and inclusions in gypsum
crystals. Insects are found as trace fossils as well
as body fossils. Fossil insects have been used in
palaeoenvironmental reconstruction in addition to
phylogenetic analysis, especially in Quaternary
deposits.
Just as there are millions of insects to be described
in today’s hot countries, so there are thousands of
fossil species to be described in places that were
once warmer. The process of collection and docu-
mentation of fossil insects is of scientific as well as
cultural value. It is possible to find more new fossil
insect species on one field trip than in an entire life-
time of collecting of the more popular fossil groups
(e.g., vertebrates). Fossil insects are thus ideal for
satisfying the goal of finding something new (and
they usually require less storage space on account of
their small size). Knowledge of the pre-Quaternary
insects has largely been forgotten since the era of
the pioneer Victorian geologists and naturalists.
There is now, however, a revival of interest, the
wider search for early (Carboniferous pre-Namurian)
 FOSSIL INVERTEBRATES/Insects 299

Figure 9 Orders through time and key events. Dashed line represents extrapolation, dots represent equilibrium (saturation) value.
Key: Pl, Pliocene; M, Miocene; O, Oligocene; E, Eocene; P, Paleocene; K, Cretaceous; J, Jurassic; Tr, Triassic; P, Permian; C,
Carboniferous; D, Devonian. Numerals 1, 2, and 3 represent Lower, Middle, and Upper subperiods and epochs; I, earliest hexapods;
II, earliest pterygotes; III, near modern range of plant-feeding strategies; IV, evolution of holometabolous insects; V, earliest insectan
parasites and parasitoid radiation; VI, earliest amberized and definite social insects.

Table 2 Principal groups of insects in the fossil record

Principal group Order Age/description

Apterygota Collembola (springtails) Lower Devonian–Recent

Diplura (two-pronged bristletails) Upper Carboniferous–Recent
‘Thysanura’ (three-tailed bristletails) This order is split into two groups: Archaeognatha
(Middle Devonian?–Recent) and Zygentoma (Upper
Carboniferous–Recent; e.g., silverfish). The extinct
order Monura (Upper Carboniferous–Permian) with
single ‘tails’ is probably related to the Archaeognatha,
whereas Zygentoma is related to the Pterygota
Pterygota
Paleoptera Palaeodictyopteroid group An Upper Carboniferous–Upper Permian group of three or
four extinct orders (Palaeodictyoptera,
Permothemistida, Megasecoptera, Diaphanopterodea)
with beak-like mouthparts and an additional pair of
‘winglets’
Ephemeroptera (mayflies) Upper Carboniferous–Recent
Protodonata Upper Carboniferous–Triassic, including the giant
dragonflies with wingspans up to 70 cm, the largest
insects of all time
Odonata (dragonflies, damselflies) Upper Carboniferous–Recent
Neoptera ‘Protorthoptera’ Lower Carboniferous–Triassic. A taxonomic wastebasket
of early neopterans

Continued
300 FOSSIL INVERTEBRATES/Insects
Table 2 Continued
Principal group Order
Polyneoptera Plecoptera (stoneflies)
Embioptera (web-spinners)
Phasmatodea (stick insects)
Orthoptera (crickets, grasshoppers,
katydids, locusts)
Titanoptera
Grylloblattodea (ice bugs)
Mantophasmatodea
Protelytroptera
Dermaptera (earwigs)
Miomoptera
Blattodea (cockroaches)
Isoptera (termites)
Mantodea (praying mantises)
Caloneurodea
Paraneoptera Zoraptera (angel insects)
Psocoptera (bark and book lice)
Phthiraptera (lice)
Thysanoptera (thrips)
Hemiptera (true bugs)
Holometabola Glosselytrodea
(Oligoneoptera) Strepsiptera (stylopids)
Coleoptera (beetles)
Raphidioptera (snake flies)
Megaloptera (alder flies)
Neuroptera (lacewings)
Hymenoptera (wasps, ants, bees)
Trichoptera (caddis flies)
Lepidoptera (moths, butterflies)
Diptera (true flies)
Siphonaptera (fleas)
Mecoptera (scorpionflies)
insects being of paramount importance. For logistic
reasons, the study of fossil insects (palaeoento-
mology) relies on international co-operation; to fac-
ilitate this objective and to promote knowledge, the
International Palaeoentomological Society was
founded in 2001.
See Also
Fossil Invertebrates: Arthropods.
Further Reading
In addition to the print literature, several groups have
web sites that are sources of information about insects:
the Arthropod Laboratory of the Russian Academy of
Sciences (http://www.palaeoentomolog.ru), the Inter-
national Palaeoentomological Society (http://www.cwru.
Age/description
Lower Permian–Recent
Lower Permian–Recent
Upper Permian–Recent
Upper Carboniferous–Recent
Extinct Triassic order allied to Orthoptera
Lower Permian–Recent
Eocene–Recent
Extinct Permian earwiglike insects
Lower Jurassic–Recent
Extinct Upper Carboniferous–Lower Jurassic insects
Upper Carboniferous–Recent
Lower Cretaceous–Recent
Lower Cretaceous–Recent
Extinct Upper Carboniferous–Permian insects
Oligocene–Recent
Lower Permian–Recent
Eocene–Recent
Lower Permian–Recent
Upper Carboniferous–Recent
Extinct Lower Permian–Upper Jurassic insects
Eocene–Recent
Lower Permian–Recent
Upper Permian–Recent
Lower Permian–Recent
Lower Permian–Recent
Upper Triassic–Recent
Lower Permian–Recent
Lower Jurassic–Recent
Lower Triassic–Recent
Lower Cretaceous–Recent
Lower Permian–Recent
edu/affil/fossilinsects), and the University of Barcelona’s
Meganeura Palaeoentomological Newsletter (http://
www.ub.es/dpep/meganeura/meganeura.htm).
Benton MJ (ed.) (1993) The Fossil Record 2. London:
Chapman & Hall.
Carpenter FM (1992) Superclass Hexapoda. Treatise on
Invertebrate Paleontology, Part R, Arthropoda 4, 3 & 4.
Earl of Cranbrook (1996) The scientific value of collec-
tions. Sarawak Museum Journal 50(71): 73–86.
Jarzembowski EA (2001) Insect ‘‘bioerosion’’. Acta Geolo-
gica Leopoldensia 26(52/53): 161–164.
Jarzembowski EA (2003) Palaeoentomology: towards the
big picture. Acta Zoologica Cracoviensia, Krakow
46(suppl.): 25–36.
Jarzembowski EA and Ross A (1993) The geological record
of insects. Geology Today 9(6): 218–223.
Rasnitsyn AP and Quicke DLJ (eds.) (2002) History of
Insects. Dordrecht: Kluwer Academic Publishers.
Wilson EO (1992) The Diversity of Life. Cambridge, MA:
Harvard University Press.