274 FOSSIL INVERTEBRATES/Arthropods

FOSSIL INVERTEBRATES

Contents

Arthropods
Trilobites
Insects
Brachiopods
Bryozoans
Corals and Other Cnidaria
Echinoderms (Other Than Echinoids)
Crinoids
Echinoids
Graptolites
Molluscs Overview
Bivalves
Gastropods
Cephalopods (Other Than Ammonites)
Ammonites
Porifera

depending on the exact nature of the leg appendages.

Arthropods
L I Anderson, National Museums of Scotland,
Edinburgh, UK
ß 2005, Elsevier Ltd. All Rights Reserved.
Introduction
Arthropods are the most diverse group of fossil and
living invertebrate animals. They have colonized
land, sea, and air due to a relatively plastic and
adaptable body plan organization. Many different
phyla of metazoans, including the arthropods, show
repetition of structure along their long axis. This is
generally termed metamerism.
Arthropod bodies are characteristically segmented,
and the different major groups are defined by various
patterns of tagmosis (segment fusion) to form heads,
bodies, and tails. The phylum derives its name from
the characteristic jointed appendages found in all
members (Greek: aryros ‘arthros’, jointed; podos
‘podos’, foot). These jointed appendages, which con-
sist of individual elements called podomeres, take
the form of walking legs, gill branches, or antennae.
Broad divisions can be made within the arthropods
Uniramians, or arthropods with single-branched ap-
pendages, retain only a walking or swimming leg.
Biramians have an appendage which consists of two
separate elements: a leg branch for movement and a
gill branch used in respiration.
Arthropods all possess a rigid exoskeleton which
provides support and protection, and acts as a buffer
between the external environment and internal body
processes. The exoskeleton of arthropods is com-
posed of a carbohydrate polymer called chitin. In
some arthropod groups, the chitin superstructure is
reinforced with mineral salts, such as calcium carbon-
ate or calcium phosphate. Arthropods with this bio-
mineralization show a higher preservation potential
than non-mineralized forms, and consequently have a
more complete fossil record through time. For non-
mineralized forms, we have to look to sites of excep-
tional preservation (Fossil Konservat-Lagerstätten) to
trace their evolutionary history and discover aspects
of their palaeobiology. Localities, such as the Middle
Cambrian Burgess Shale, the Ordovician Soom
Shale, the Devonian Rhynie chert, the Carboniferous
Mazon Creek nodule hosted biota, the Jurassic Soln-
hofen Lithographic limestone, and Baltic, Domin-
ican, and Mexican ambers, all provide windows
onto the arthropod faunas at stratigraphical points

in geological time in which preservational failure
would otherwise obscure our view of evolution of
the various groups (see Lagerstätten). Furthermore,
there is a gradation of fidelity of preservation even
in these special examples. To this end, it is entirely
possible to study the exquisite three-dimensional
outer surfaces of Tertiary flies in amber (Figure 1) or
the internalized respiratory systems of Devonian
arachnids.
A consequence of the possession of a rigid exoskel-
eton composed of cuticle is that, in order for the
arthropod to increase in body size over time, the
rigid cuticle has to be shed or moulted and a new
cuticle grown below the surface to replace it. Arthro-
pods differ from many other animal groups in that
they go through the process of ecdysis (literally shed-
ding the exoskeleton as a moult) in order to attain
greater body size. Ecdysis is facilitated by the pres-
ence of sutures in the exoskeleton, lines of weakness
which can be broken through so that the arthropod
can struggle free of the old skin. The ontogeny of
Figure 1 A fungus gnat preserved in Baltic amber from
the Upper Oligocene–Lower Eocene of Palanga, Lithuania
(
55). ß The Trustees of the National Museums of Scotland,
Edinburgh, UK (NMS G.2002.6.17).
FOSSIL INVERTEBRATES/Arthropods 275
ecdysis allows segment addition and tagmosis to be
identified from the first instars right up to the adult
forms. Unfortunately, for arthropods, moulting of the
old skin and the time taken to harden the new exo-
skeleton leaves them vulnerable to predator attack or
adverse environmental conditions. For this reason,
many arthropods (both in aquatic and terrestrial en-
vironments) seek out a ‘moulting refugia’ in which
the process of ecdysis and subsequent hardening of
the new exoskeleton can take place in relative safety.
Where mortality occurs during this post-moult stage,
the cuticle of the arthropod fossil formed can display
wrinkles, creases, or even tears in the structure.
The cuticle itself takes the form of hardened scler-
ites, one per body segment. Dorsal sclerites are termed
tergites, ventral sclerites are termed sternites. Where
the tergites and sternites are fused together, they form
a simple ring structure enclosing the arthropod soft
tissues. Lying between these discrete segments are
areas of connective and unmineralized cuticle. The
presence of this less robust construction material ex-
plains why, upon death, well-mineralized arthropods,
such as trilobites, can disarticulate to a greater or
lesser degree into their constituent parts (cephalon,
thorax, thoracic segments, pygidium, etc.). The leg
appendages in arthropods are similarly constructed
with cylindrical reinforced cuticular podomeres with
intervening, less well-sclerotized arthrodial mem-
branes connecting them. Muscle attachments re-
quired to move the individual leg podomeres are
attached to small boss-like projections from the inside
of the exoskeleton, called apodemes.
The cuticle of arthropods explains in part their
success in the colonization of such a wide variety
of different environments with widely varying chal-
lenges. The cuticle provides a physical barrier between
the internal organs and the external environment
(which helps to reduce desiccation through water
loss in terrestrial forms) and a chemical barrier in
aquatic conditions (where hypo- or hypersalinity
could upset the osmotic balance, leading to dehydra-
tion of the organism and ultimately death). As well as
a physical barrier, waxes produced by the cuticle layer
help to waterproof this material. Some other more
unusual properties of arthropod cuticle have been
identified recently, including the fluorescence under
ultraviolet (UV) light of ‘hyaline’ cuticle within the
exoskeleton of scorpions. It happens that this ‘hyaline’
layer is actually very resistant to bacterial degrad-
ation and is therefore often the main remaining con-
stituent material of fossil scorpions. For example,
the Early Carboniferous East Kirkton biota of West
Lothian, Scotland, contained a number of different
scorpion taxa which were recovered by hydrofluoric
acid maceration of the enclosing limestone matrix.
276 FOSSIL INVERTEBRATES/Arthropods
The level of morphological detail made available
through this technique was as good as studying extant
species.
Classification Schemes
There still remains scope for significant disagreement
in the higher level taxonomy of this group of animals.
Historically, the main dichotomy relates to whether
the phylum Arthropoda is a monophyletic or poly-
phyletic group of organisms. Some variation occurs in
the higher level classification of this group, with some
authors recognizing Arthropoda at phylum level and
the constituent members variously as subphyla or
orders. A monophyletic grouping for the arthropods
would imply that all of the lower level groupings
originated from a single common ancestor. This was
the largely historic viewpoint based on the striking
common appearance of the various groups. In order
to counter what seems to be the logical conclusion
based on many shared characters in common, the
argument for polyphyly has to promote plausible
explanations for the observed similarities between
groups as well as providing proof of separate points
of origin. The fossil record may provide just such
evidence, allowing the evolution of the various
arthropods to be considered with the benefit of deep
time hindsight. However, since the work of Sidnie
Manton, who suggested that three major groups, the
Crustacea, the Chelicerata, and the Uniramia, would
more comfortably be accommodated as phyla of their
own, the consensus view has swung back to the like-
lihood of the monophyletic status of the group being
the correct conclusion. This has been due in part to
the advent of molecular sequencing techniques. Novel
(non-morphological) hypotheses linking crustaceans
and hexapods were primarily based on the analysis of
single or a handful of genes. However, as the science
advances and the techniques of extraction improve,
the case for monophyly appears to be more solid.
Indeed, one study not only supports the crustacean–
hexapod clade, but also recognizes pycnogonids as
the sister group to other euarthropods. The historical
convention has been to subdivide the Arthropoda into
the following equal-biased groupings:
. Trilobita (containing all trilobites, but excluding
naroiid arthropods);
. Chelicerata (Arachnida, Scorpionida, Xiphosura,
Eurypterida, Chasmataspida);
. Crustacea;
. Uniramia (including both insects and myriapods).
However, as more and more examples of basal
arthropods and ‘oddball’ taxa are discovered, primar-
ily from Fossil Konservat-Lagerstätten, other less
conventional higher taxon groupings can be identified.
Some examples include:
. Atelocerata (Diplopoda, Chilopoda, Pauropoda,
Symphyla, Hexapoda);
. Arachnomorpha/Arachnata (Trilobites, Chelicerates,
Trilobitomorpha).
Arachnomorpha is a clade which basically consists
of all taxa more closely related to Chelicerates than
crustaceans. Panarthropoda is again a clade, but
this time containing all true arthropods and their
soft-bodied relatives, such as onychophorans and
tardigrades.
To this end, the term ‘arthropod’ should probably
be used to represent a grade of organization in which
a soft-bodied ancestor (presumably a segmented
worm) developed a toughened cuticle. Gene sequen-
cing and molecular markers may provide fascinating
new insights into the interrelationships of the various
high-level arthropod groups.
What the Fossil Record Says
Tracing arthropods back through geological time in-
dicates that the divisions between the major groups
extend almost as far back as the fossil record of
Metazoa. Convergence of form and channelling by
functional morphology of multiple arthropodization
events through forming a rigid cuticle could have led
to polyphyly; however, the more complete our record
of the group becomes, the less likely this seems.
A Brief History
The first arthropods appear in the fossil record of life
on Earth in the Lower Cambrian some 540 Ma. The
arthropod faunas of the Chengjiang biota of China
demonstrate that, even at this early stage, a diverse
range of arthropod body plans had evolved. By
Middle Cambrian times, rapid and explosive radi-
ation and evolution of arthropods led to the devel-
opment of many different body plans, including
the majority of those still extant. This ‘Cambrian
Explosion’ in the number and diversity of arthropo-
dan groundplans was one of the first major radiations
of the group and took place in the marine envir-
onment. By the Upper Cambrian, trilobites had di-
versified markedly and this carried through into
Ordovician times.
Mid-Silurian and Lower Devonian rocks yield
evidence of the next major step in the evolution of
the arthropods: the colonization of the terrestrial
environment, a process termed terrestrialization. Dif-
ferent terrestrial arthropod groups probably took
different terrestrialization pathways. Although this
 FOSSIL INVERTEBRATES/Arthropods 277

is difficult to determine from the fossil forms, extant within the Crustacea continues to the present day,
members of the ancient lineages can lend clues with arthropods still developing new and successive
as to what these pathways might have been. For waves of colonization, particularly in terms of the
example, osmotic concentrations of body fluids in terrestrialization of some decapod crustaceans, such
myriapods suggest that this group terrestrialized as the land crabs.
straight from the sea without an intermediate step in
freshwater.
Respiration
One particular locality which has yielded a dispro-
portionately large amount of data on early terres- From the same basic body plans, different solutions to
trial ecosystems is the Devonian Rhynie Chert the challenges of respiration in aquatic and terrestrial
Lagerstätte in Aberdeenshire, Scotland. Here, the environments have evolved. In the smallest of micro-
silica-mineralizing action of hot spring and geyser arthropods, cutaneous respiration, by which the skin
fluids captured, in three-dimensional detail, early acts as a gas exchange surface, takes place. The ratio
terrestrial arthropods, such as arachnids, insects, of surface area to internal volume limits the max-
and centipedes, as well as aquatic forms, such as imum size of arthropod which can utilise this method.
freshwater crustaceans and euthycarcinoids. The re- In larger aquatic arthropods, respiratory gas ex-
markable morphological stasis in early terrestrial change is through structures termed ‘book gills’.
arthropods, in relation to extant forms, suggests that These thin lamellate cuticular structures are bathed
arthropod-dominated terrestrial ecosystems stabil- in oxygen-containing water, either wholly external to
ized early on in the Earth’s history and that soil and the body, as in the gill branches of trilobites, or in a
litter habitats and inhabitants have changed very little specialized cavity, as in the horseshoe crabs (Figure 2).
ever since. Terrestrial arthropods show adaptation to the par-
One of the novel innovations of arthropods was ticular challenges of living in an environment in
the evolution of muscle-powered flight in the insects which the constant danger of desiccation means that
(see Fossil Invertebrates: Insects). The earliest insects
or apterygotes (insects without wings) first appear in
sediments of Devonian age in the form of collembo-
lans and thysanurans. In earliest Carboniferous times,
the major expansion and evolution of land-based
flora provided new ecological niches available to
this highly adaptable group. The advent of insects
and the development of flight led to the colonization
of an entirely new ecospace. Interestingly, the earliest
non-scorpion arachnids appear to have relied solely
on poisoned fangs to capture and subdue prey items.
Orb web-weaving using spider silk could only have
developed in response to the presence of aerial insects.
By Lower Carboniferous times, the first winged
insects had developed and an explosion in the diver-
sity of insects occurred due to coal swamp forest
ecosystems. Preservation and discovery are aided by
the presence of siderite concretions in the roof shales
to many coal seams which, in turn, are commercially
exploitable. During the Middle Carboniferous, in-
sects became diverse and widespread for the first
time in their fossil record. This trend continued
through the Carboniferous and into the Permian
period, whereupon an explosion in the total diversity
of fossil insects took place. The depositional environ-
ments in which these fossils are found do not differ
significantly, suggesting a real increase in diversity.
During the Jurassic and Cretaceous, favourable shal-
Figure 2 The xiphosuran, Rolfeia sp., from the Lower
low (epieric) sea conditions saw the emergence and Carboniferous of Mumbie Quarry, Glencartholm, Dumfries and
dominance of decapod crustaceans as a major player Galloway, Scotland (
1.1). ß The Trustees of the National
in nearshore marine ecosystems. This development Museums of Scotland, Edinburgh, UK (NMS G.1998.35.3).
278 FOSSIL INVERTEBRATES/Arthropods
book gills would cease to function. In terrestrial
arthropods, respiration is via book lungs (analogous
and probably derived in part from book gills) or by a
tracheal system. The lamellate structure of book
lungs requires additional supporting struts so that
they do not collapse in air. Furthermore, the book
lungs tend to be positioned internally within the
body of the animal as a means of further reducing
water loss through desiccation. The tracheal system
is a passive method of diffusing oxygen into the body
of the arthropod via a branching system of tubes
opening on the surface of the exoskeleton at a tra-
cheole. The insects show the greatest development of
the tracheal system, related no doubt to the oxygen
demands of muscle-powered flight. Other unira-
mians, such as millipedes, also respire in a similar
manner through the presence of a series of paired
spiracles along the length of the body. The earliest
example of this form of respiration is recorded in
the mid-Silurian millipede, Pneumodesmus newmani,
from Stonehaven, Scotland.
The diffusive process involved in tracheal respir-
ation places an upper limit on the maximum size of
terrestrial arthropods which utilize this method,
unless a means of secondary ‘pumping’ derived from
muscle contraction can improve efficiency. Book
gills, on the other hand, operate in water and are
not subject to the same constraints on size which are
faced by a land-based book lung system. In some
respects, this may help to explain why the largest
arthropod fossils found are those of aquatic organ-
isms, such as the extinct eurypterids. Indeed, some
very active arachnids, such as solfugids (camel spi-
ders), possess both book lung and tracheal systems,
allowing them to meet the oxygen demands of their
highly active and voracious life style. The antiquity
of the development of tracheal respiratory systems
in terrestrial arthropods stretches back as far as
the earliest opilionid (harvestman spider) found in
the Early Devonian Rhynie Chert, Aberdeenshire,
Scotland.
Reproduction
Aquatic and terrestrial environments present differ-
ent challenges to reproductive strategies in arthro-
pods. Terrestrial arachnids tackled the problem of
the desiccation of reproductive material by producing
a spermatophore, which is either passed from male to
female using specially adapted leg appendages or
pedipalps (spiders), or left on the ground attached to
a stalk (scorpions).
Internal fertilization is practised by a few groups,
including the opilionids (harvestman spiders), and the
preservation of internalized organs in arachnids, such
as the Rhynie Chert opilionids, in which penis and
ovipositor structures are known in separate specimens,
demonstrates morphological conservatism.
Direct morphological evidence of sexual di-
morphism, or reproductive structures, has long been
identified in the chelicerate arthropod group, the
eurypterids. In these arthropods, Type A and Type B
appendages have been identified on the ventral sur-
faces. A recent study went further and demonstrated
that the longer Type A appendage was probably
female, whilst the Type B appendage belonged to the
male. An elaborate system of flexure along predefined
sutures in the cuticle allowed the female to retrieve an
unstalked spermatophore from the substrate prior to
storage in the spermathecae.
Feeding
An effective demonstration of how the understanding
of the form and feeding function of extant arthropods
could be ‘reverse engineered’ to understand those of
the fossil forms was provided by one particular
example: the Lipostracan crustacean, Lepidocaris,
known from three-dimensional fossils in the Early
Devonian Rhynie Chert.
The most recent synthesis of feeding in trilobites
(see Fossil Invertebrates: Trilobites) suggested that, as
a group, the various trilobites employed different
strategies. Indeed, it was argued that much of the
variety of form seen in the exoskeletal structure of
trilobites was probably driven in response to specific
feeding modes (Figure 3). In the main, the structures
most responsive to this driving factor were the cepha-
lon (or head region) and the hypostome, a small plate
lying anterior to the mouth cavity and variously free
or fixed depending on the feeding strategy. The primi-
tive mode of feeding in trilobites was identified as
predatory or scavenging, with cephalon and accom-
panying hypostome morphology (coupled with over-
all body shape) used as a guide to indicate possible
feeding strategies within different trilobite groups.
However, this may even be a relict from the
arthropods ancestral to the trilobites.
On the other hand, detritivorous trilobites pos-
sessed a hypostome detached from the doublure of
the cephalon. The glabella (the axially aligned raised
area atop the cephalon), where the stomach was
thought to reside, would be relatively small in these
forms, reflecting the small quantities of food pro-
cessed at any one time. Filter-feeding trilobites exhibit
a different overall morphology again, and often pos-
sess a highly vaulted cephalon which would have
allowed the leg appendages to stir into suspension
food-bearing sediment particles. The demonstration
of an elegant scheme such as this in a reasonably

Figure 3 The trilobite, Phacops sp., from the Devonian of Mo-
rocco, North Africa (
1.7). ß The Trustees of the National
Museums of Scotland, Edinburgh, UK (NMS G.1987.20.4).
well-known fossil group should be applied to the
less well-known Palaeozoic arthropods.
Relevance
Arthropod fossils are used by palaeontologists and
geologists in a number of different ways for different
reasons. In rocks of Palaeozoic age, the trilobites have
long been used as biostratigraphical marker fossils.
They find application in this particular field of study
as they are readily fossilized (carbonate-impregnated
exoskeleton), numerically abundant (each individual
moults many times over a lifetime, ‘amplifying’ their
number of potential fossils), and show sufficiently
rapid speciation events to make them useful as index
fossils. In Early Palaeozoic rocks, nektonic species
are useful in palaeogeographical reconstructions of
former oceans, seas, and, by default, landmasses; the
nektonic life style has led to a widespread occurrence
of the resultant fossil remains, ideal for broad
stratigraphical correlations. Another use for the
numerically abundant trilobites in a stratigraphical
framework is the evaluation of evolutionary rates
within well-defined lineages. Such frameworks allow
trilobites to be utilized in studies of evolutionary
FOSSIL INVERTEBRATES/Arthropods 279
changes within a species, up until the point at which
a new species evolves, a field of study known as
micro-evolution.
Finally, many micro-palaeontological samples,
such as fossils of ostracodes (see Microfossils: Ostra-
coda), are similarly numerically abundant and readily
preserved, and have found application in palaeonto-
logical and archaeological studies of past environ-
ments due to their environmental sensitivity. The
calcium carbonate-impregnated shells of these small
bivalve-carapaced crustaceans are a common constit-
uent in rocks of all ages from the Cambrian onwards.
Visual Systems
Two main types of eyes are recognized in the arthro-
pods: compound (multifaceted) and simple ocellar
eyes (Figure 4). Compound eyes probably developed
separately in a number of different arthropod groups,
giving rise to analogous if not homologous structure.
Many arthropods possess both compound and simple
eyes, as demonstrated by the extant chelicerate horse-
shoe crab, Limulus. The ocellar eyes are positioned
on the top of the dorsal surface of the carapace and
are primarily used to detect light levels, indicating
whether or not the top of the carapace is covered
by sand; this information is of particular use to
burrowing arthropods. The compound eyes of trilo-
bites are further subdivided into holochroal and schi-
zochroal forms, with the schizochroal structure
developing as a result of paedomorphosis. Holo-
chroal trilobite eyes consist of many round or polyg-
onal lenses, the edges of which are all in contact and
covered by a single corneal membrane. Schizochroal
eyes are a unique system restricted to certain trilobite
groups (Phacopina). The lenses are large and separ-
ated from one another. Each separate lens has its own
separate corneal covering. Mounting the eye lens on
upstanding and curved turrets may have allowed
certain trilobite species a full 360 field of vision.
Trace Fossils
The combination of rigid appendages and the ability
to be motile in a wide range of ecological habitats has
provided a rich and diverse trace fossil record which
can be assigned to the activities of arthropods. Trace
fossils can indicate the presence of arthropods in de-
positional environments in which preservation would
normally fail, and body fossils are non-existent.
A good example of this is the Permian Coconino
Sandstone Formation of Arizona. Various studies of
the desert sandstones have revealed the presence of
scorpion and tarantula trackways, the body fossils
of which are absent from these environments.
280 FOSSIL INVERTEBRATES/Arthropods
Figure 4 The enigmatic arthropod, Ainiktozoon loganense, from the Silurian Fossil Konservat-Lagerstatten of Lesmahagow, Lanark-
shire, Scotland (
1.5). ß The Trustees of the National Museums of Scotland, Edinburgh, UK (NMS G.1996.40.1).
Distinctive trace fossils, such as Kouphichnium,
can reveal the presence of xiphosuran arthropods
(horseshoe crabs) and, by analogy with their extant
relatives, such as Limulus, can be used to imply near-
shore or freshwater palaeoenvironments in otherwise
sediments devoid of body fossils. Trace fossils, when
found in conjunction with body fossil assemblages,
may allow behaviour to be interpreted, such as the
speed of movement or different actions (nesting, for-
aging, etc.). Of particular importance in the fossil
record of insects is the formation of leaf mines and
bite marks in accompanying floral assemblages,
giving a clear indication of an escalating arms race
between plants and arthropods.
Final Remarks
In conclusion, the Arthropoda can be marked out as
one of the most important groups of organisms on the
Earth. Their fossil record spans from the earliest
recognizable invertebrate faunas right up until the
present day. Their numerical abundance, both in the
present and in the past, positions them as the major
players in ecological structure: predators and prey in
all three major Earth environments (land, sea, and
air). The group provides excellent opportunities to
investigate the effects of extinction events, adap-
tive radiation, macro- and micro-evolution, and the
consequences of palaeobiology in extinct groups.
Understanding the relationships between arthropods
and other plant and animal groups remains import-
ant. Arthropods were an important part of the food
chain in the past, just as they are today. Further
advances in DNA sequencing and other types of mo-
lecular data will no doubt increase our knowledge of
the inter-relationships of this diverse and fascinating
group.
See Also
Evolution. Fossil Invertebrates: Trilobites; Insects. La-
gerstätten. Microfossils: Ostracoda. Palaeoecology.
Trace Fossils.
Further Reading
Allen KC and Briggs DEG (1989) Evolution and the Fossil
Record. London: Belhaven Press, Pinter Publishers.
Benton MJ (ed.) (1993) The Fossil Record 2. London:
Chapman and Hall.
Braddy SJ and Dunlop JA (1997) The functional
morphology of mating in the Silurian eurypterid,
Baltoeurypterus tetragonophthalmus (Fischer, 1839).
The Zoological Journal of the Linnean Society 121:
435–461.
Briggs DEG and Crowther PR (eds.) (1990) Palaeobiology:
A Synthesis. Cambridge: Cambridge University Press.
Edgecombe DG, Wilson GDF, Colgan DJ, Gray MR, and
Cassis G (2000) Arthropod Cladistics: combined analysis
of histone H3 and u2 snRNA sequences and morphology.
Cladistics 16: 155–203.
 FOSSIL INVERTEBRATES/Trilobites 281

Fortey RA and Owens RM (1999) Feeding habits in trilo-
bites. Palaeontology 43: 429–465.
Fryer G (1985) Structure and habits of living branchiopod
crustaceans and their bearing on the interpretation
of fossil forms. Transactions of the Royal Society of
Edinburgh: Earth Sciences 76: 103–113.
Rasnitsyn AP and Quicke DLJ (2002) History of Insects.
Dordrecht, Boston, London: Kluwer Academic
Publishers.
Willmer P (1993) Invertebrate Relationships: Patterns in
Animal Evolution. Cambridge: Cambridge University
Press.

Trilobites
A W A Rushton, The Natural History Museum, that more than 5000 genera have now been de-
London, UK scribed and a huge number of species named, perhaps
Copyright 2005, Natural History Museum. All Rights Reserved. approaching 20 000.

Introduction Form of the Exoskeleton

The general features of the exoskeleton are labelled in
The trilobites are a large group of extinct marine
Figure 2. The exoskeleton typically has an oval out-
arthropods, the Class Trilobita, characterized by the
line in plan view, and consists of a cephalon (head-
longitudinal division of the exoskeleton into three
shield), furnished with a pair of compound eyes, a
lobes – a convex central (axial) lobe flanked on each
jointed thorax of few to many segments, and a pygi-
side by a flatter pleural area (Figure 1) – hence their
dium (tail-piece) made up of fused segments. Typic-
name. Trilobites were widely distributed throughout
ally the animal was 2–10 cm in length, but a species of
the Palaeozoic era, first appearing in the later part
Acanthopleurella is just over 1 mm in length when
of the Early Cambrian period (at about 520 Ma)
fully grown, and the largest known Isotelus is 72 cm
and disappearing just prior to the end-Permian mass
in length.
extinction (ca. 250 Ma). During their 270 million
The trilobite exoskeleton, unlike that of many art-
year existence, trilobites spread worldwide, occupied
hropods, was mineralized by calcite and phosphate,
a wide range of marine habitats, and evolved a great
thus greatly increasing its potential for preservation
variety of morphologies.
Trilobites are a relatively well-studied group and
a good deal is known, or inferred, about their life,
growth, and activities. Their extensive fossil record
has provided illuminating case histories in evolution
and ecological adaptation, and, in addition, they have
proved to be of great value in stratigraphy, especially
in the Cambrian and Ordovician periods, and in
interpreting Palaeozoic palaeobiogeography.
Thanks to their striking appearance, strange-looking
yet evidently organic in origin, trilobites have always
been attractive fossils. Specimens have been recorded
in prehistoric burial sites and they have found a place
in folk-lore; in more recent times, avid collectors have
built up magnificent collections of fine specimens,
some of which have commanded high prices. The
scientific study of trilobites commenced at the end
of the seventeenth century, and, during the eight-
eenth and nineteenth centuries, European workers,
including Linnaeus, published descriptions and illus-
trations. The benchmark was Barrande’s illustrated
account of the Bohemian trilobites, published in
1852, that showed their astonishing variety and Figure 1 The dorsal exoskeleton of Neometacryphaeus (sub-
stimulated interest in the group amongst all scientific order Phacopina) from the Devonian of Morocco. About natural
communities; this interest continues to this day, such size. Courtesy of Professor R. A. Fortey.