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Jarzembowski2005 PDF
Jarzembowski2005 PDF
Insects
E A Jarzembowski, University of Reading, Reading, How many insects are there? Nobody knows
UK and Maidstone Museum and Bentlif Art Gallery, for sure, although all estimates suggest that millions
Maidstone, UK of insect species remain to be described. The great
ß 2005, Elsevier Ltd. All Rights Reserved. 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
Introduction
Insects, which belong to the taxonomic group
Hexapoda (‘six legs’; Table 1), are the most successful Table 1 Insects – systematic position
organisms on Earth, if biodiversity is measured as Taxonomic division Group/number
a count of the number of species. More than
Kingdom Animalia
1.4 million species of insects have been described in
Phylum Arthropoda
the past 250 years, comprising 65% of all known Superclass Hexapoda (insects in the
species of life on Earth (Figure 1A). Insects are also broad sense)
the most successful group in the fossil record, if Order About 43
palaeodiversity is measured as a count of the number Family Over 1500
Genus ?
of families documented, as many palaeontologists
Species 3–20 million
do (Figure 2).
296 FOSSIL INVERTEBRATES/Insects
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-
Figure 1 Global biodiversity, showing percentage breakdown
sities (100 000 or more species) are reached in only
for insects and other organisms. (A) Percentages of the app- four (holometabolous) orders: Coleoptera (beetles),
roximate number of known species worldwide (1 454 000). Lepidoptera (moths and butterflies), Hymenoptera
(B) Percentages of the estimated total number of species (wasps, ants, and bees), and Diptera (true flies).
worldwide, which includes those thought to be undiscovered
(65 654 000).
Geological History
The origin of insects is a mystery, the Cambrian
development. Simple extrapolation from the fossil Burgess Shale arthropods being too early to cast any
record suggests that the total number of insect species light on the subject. The oldest definite hexapod is
is probably less than 20 million, although some biolo- Rhyniella praecursor from the Early Devonian Rhynie
gists prefer a higher figure. Everyone is agreed, never- Chert. Rhyniella praecursor is a springtail belonging
theless, that insects represent over 50% of all known to the living apterygote order Collembola. The earli-
species and that they belong to an exclusive group of est true insect is currently considered to be Rhyniog-
hyperdiverse organisms. 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-
Origins ing paleopterans and polyneopterans. These insects
Where do insects come from? Insects are undoub- were the world’s first flying animals, long before ver-
tedly a class of arthropods, or ‘joint-legged’ animals tebrates took to the air. In the succeeding Permian, the
(Table 1). Arthropods (see Fossil Invertebrates: paraneopterans and holometabolans became estab-
Arthropods) also include trilobites, crustaceans lished. Insects are essentially terrestrial organisms,
(prawns, etc.), chelicerates (spiders, etc.), and myria- but the first definite freshwater forms appeared in
pods (millipedes and centipedes). Insects show closest the Permian. After a setback in the Early Triassic
relationships to crustaceans (e.g., in the structure of extinction (Figure 8), insects regained their ordinal
their compound eyes) and myriapods (e.g., in their strength by the Tertiary, establishing some new innov-
tubular or tracheate respiratory system) (Figure 3). ations on the way, e.g., evolving parasitic and para-
The exact relationships are currently a subject of sitoid forms as well as insect societies (Figure 9). The
debate. 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 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.
Continued
300 FOSSIL INVERTEBRATES/Insects
Table 2 Continued
insects being of paramount importance. For logistic edu/affil/fossilinsects), and the University of Barcelona’s
reasons, the study of fossil insects (palaeoento- Meganeura Palaeoentomological Newsletter (http://
mology) relies on international co-operation; to fac- www.ub.es/dpep/meganeura/meganeura.htm).
ilitate this objective and to promote knowledge, the Benton MJ (ed.) (1993) The Fossil Record 2. London:
Chapman & Hall.
International Palaeoentomological Society was
Carpenter FM (1992) Superclass Hexapoda. Treatise on
founded in 2001.
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.
See Also Jarzembowski EA (2001) Insect ‘‘bioerosion’’. Acta Geolo-
Fossil Invertebrates: Arthropods. gica Leopoldensia 26(52/53): 161–164.
Jarzembowski EA (2003) Palaeoentomology: towards the
big picture. Acta Zoologica Cracoviensia, Krakow
Further Reading 46(suppl.): 25–36.
Jarzembowski EA and Ross A (1993) The geological record
In addition to the print literature, several groups have of insects. Geology Today 9(6): 218–223.
web sites that are sources of information about insects: Rasnitsyn AP and Quicke DLJ (eds.) (2002) History of
the Arthropod Laboratory of the Russian Academy of Insects. Dordrecht: Kluwer Academic Publishers.
Sciences (http://www.palaeoentomolog.ru), the Inter- Wilson EO (1992) The Diversity of Life. Cambridge, MA:
national Palaeoentomological Society (http://www.cwru. Harvard University Press.