Arthropoda (Arthropods)
Neil W Blackstone, Northern Illinois University, De Kalb, Illinois, USA
The Arthropoda is a phylum of ecdysozoan animals in which the epidermis produces a
segmented, jointed and hardened exoskeleton, which has internal musculature.
Basic Design
Arthropods generally exhibit ecdysozoan features such as
the presence of a cuticle, reduction or absence of locomotory
cilia and a cuticular moulting cycle regulated by ecdyster-
oids. Arthropods (or panarthropods) nevertheless share de-
rived morphological characteristics that distinguish them
from other ecdysozoa, for instance, compound eyes and a
segmented, jointed and hardened exoskeleton, which has
internal musculature. Arthropod segments usually arise by
teloblastic growth; each segment primitively bears a pair of
jointed limbs and exhibits other typical arthropod features
(Figure 1). Segments tend to be grouped into body regions or
tagmata and specialized in this regard. Major groups of
arthropods exhibit a characteristic tagmosis (Figure 2a,b); for
instance, insects and crustaceans have three tagmata – head,
thorax and abdomen – with a characteristic number of seg-
ments comprising the head. Subsidiary taxa of these major
groups may have a characteristic number of segments in the
thorax and abdomen as well. Considerable debate exists
concerning the homology of head segments in different
groups of arthropods. For instance, the structure of the ar-
thropod brain (Figure 2d) might suggest homologies between
head segments in chelicerates and crustaceans, but this is far
from clear. Arthropod limbs exhibit two forms: uniramous
(one branch) and biramous (two branched). Generally
chelicerates, myriapods and insects have uniramous limbs,
while crustaceans and trilobites have biramous limbs. Limbs
of major groups of arthropods also show characteristic
numbers, morphologies and functions. Antennae, mouth-
parts, chelicerae, walking legs, grasping and swimming
Dorsal longitudinal Heart
muscle Dorsoventral muscle
Haemocoel
Extrinsic
limb muscles
Exoskeleton
Intrinsic
Gut limb muscles
Ventral nerve
cord
Ventral longitudinal
muscle
Figure 1 A cross-section of a typical arthropod segment showing basic
characteristics. All arthropods are built from many such segments, which
may be modified in various ways.
ENCYCLOPEDIA OF LIFE SCIENCES & 2005, John Wiley & Sons, Ltd. www.els.net
Introductory article
Article Contents
. Basic Design
. Diversity and Abundance
. Habits and Habitats
. The Arthropod Cuticle
. Arthropod Physiology
. The Impact of Arthropods in the Modern World
. Fossil History
. Phylogenetic Controversy – One Group or Many?
. Modern Phylogeny
doi: 10.1038/npg.els.0004135
appendages are all derived from the basic arthropod limb
(Figures 1 and 2). Arthropods also include the only inverte-
brate group to evolve flight. Internal features (Figures 1 and
2) include a reduced coelom, an open circulatory system
with a dorsal, ostiate heart, a complete gut, a dorsal brain
and paired ganglionated, ventral nerve cords. See also:
Crustacea (crustaceans); Insecta (insects)
Diversity and Abundance
At least 75% of all animal species so far described belong to
the phylum Arthropoda, and millions of species, particu-
larly of insects, probably remain to be described. Insects,
spiders, scorpions, horseshoe crabs, centipedes, millipedes
and crustaceans are all arthropods; together these ar-
thropods are not only abundant and diverse, but also play
vital roles in all ecosystems. This dominance of arthropods
has continued since the earliest beginnings of animal life,
although different groups of arthropods have been pre-
dominant at different times in the history of life. Initially,
the now-extinct trilobites were the most prominent group;
later, the now largely extinct merostomates (horseshoe
crabs and water scorpions) achieved prominence. Today,
arachnids (scorpions, spiders, mites and ticks), crustaceans
(barnacles, copepods, amphipods, shrimps, lobsters and
crabs), myriapods (centipedes and millipedes) and insects
(beetles, flies, butterflies, moths, ants, wasps and bees) are
the major components of arthropod diversity. Notably, two
of these modern groups – crustaceans and particularly
insects – have evolved eusociality, a form of colonial,
group-living in which only some of the participating
individuals carry out reproductive functions. See also:
Chelicerata (arachnids, including spiders, mites and scor-
pions); Crustacea (crustaceans); Insecta (insects); Myria-
poda (including centipedes and millipedes); Trilobita
(trilobites)
1
 Arthropoda (Arthropods)

Simple eye Mouth

Compound
Gnathobase Chelicera
Carapace eye
Pedipalp

Prosoma
Walking
legs

Book gills
Chilaria
Opisthosoma Anus

Telson

(a) (b)

Heart
Ostium Midgut Intestine
Anterior
arteries Segmental ganglion

Gizzard

Mouth
Crop
(c) Brain Ventral Base of Anus
nerve gill
cord

Protocerebrum
Optic
nerve Deuterocerebrum
Nerve to Gut
chelicera First
Tritocerebrum antennary nerve

Secondary
(d) Ventral nerve cord antennary nerve

Figure 2 Dorsal (a) and ventral (b) views of Limulus polyphemus, the common horseshoe crab of Atlantic and Gulf coasts of North America. This xiphosuran
chelicerate shows many typical arthropod features both externally and internally, (c) comparison of the structure of the arthropod brain, e.g. that of
L. polyphemus and a crustacean (d) might suggest that the chelicerae of the former are homologous to the second antennae of the latter, but this is far from
clear.

Habits and Habitats
The diversity of arthropods corresponds to the diversity of
the habitats that they occupy. Remarkably, arthropods
have been enormously successful in marine, freshwater and
terrestrial environments. The exclusively marine trilobites
exploited a variety of habitats. Most were benthic, crawling
over the bottom or ploughing through the upper layers of
sediments. Larvae and even adults of some small species
may have been planktonic. Merostomates include the giant
(up to 3 m long) water scorpions, which occupied marine
and freshwater environments and may have even been
more or less terrestrial. Scorpions are among the most an-
cient terrestrial arthropods, inhabiting a variety of terres-
trial environments, particularly deserts and tropical
forests. Spiders are one of the most abundant types of ter-
restrial animals. They inhabit virtually every terrestrial
habitat as well as freshwater and intertidal marine habitats.
The ability of spiders to produce silk and fashion it into
webs and other devices has allowed them in some sense to

2

construct and modify the habitats that they occupy.
Arachnids are also unusual in that they lack compound
eyes. Pycnogonids (sea spiders) are widely distributed in
marine environments, functioning largely as slow-moving,
‘sit and wait’ predators. Myriapods are terrestrial, func-
tioning as active, aggressive – and poisonous – predators
(centipedes) or herbivores (millipedes). Insects are spec-
tacularly diverse in freshwater and particularly terrestrial
habitats, and less commonly inhabit the fringes of the
ocean itself. The success of insects is largely because of their
intimate, extensive and long-standing relationships with
angiosperms, the flowering plants – feeding on them, pol-
linating them, engaging in symbioses with them. The
capacity of many insects for flight has further opened a
range of habitats for them. Crustaceans, on the other hand,
are predominantly oceanic and are found at all depths in
every marine, brackish and freshwater habitat. Isopods
(pill bugs) are one of the few groups of crustaceans that are
truly terrestrial.
The Arthropod Cuticle
The characteristic hardened exoskeleton of arthropods in
part explains their success as a group. This exoskeleton or
cuticle is a complex layered structure secreted by the un-
derlying epidermal cells (Figure 3). While the cuticle has
been most extensively studied in decapod crustaceans and
insects, many of its features appear to be shared by all
arthropods. The outermost layer (the epicuticle) consists of
a protective layer of lipoprotein overlying the waxy layer.
The waxy layer is particularly well developed in terrestrial
insects and arachnids; it contains various long-chain
Epicuticle
Exocuticle
Calcified
endocuticle
Uncalcified
endocuticle
Epidermis
Basement membrane
Figure 3 The cuticle and epidermis of a crustacean.
Arthropoda (Arthropods)
hydrocarbons and fatty acids and serves to make the
epicuticle both relatively water-impermeable and an effec-
tive barrier against pathogens. The innermost layer of the
epicuticle consists of the cuticulin layer, which is made up
of proteins. While the epicuticle is functionally important,
it is also quite thin, consisting of only about 3% of the
exoskeleton’s total thickness.
The bulk of the cuticle consists of the procuticle, formed
from layers of protein and chitin. The procuticle is some-
times divided into an exocuticle and an endocuticle. The
procuticle is tough, but not necessarily rigid. Two pro-
cesses produce a hardened, inflexible procuticle: mineral-
ization and sclerotization. The former process involves the
deposition of calcium carbonate and is common in crus-
taceans inhabiting mineral-rich seawater. The latter pro-
cess involves ‘tanning’ of the protein component by the
formation of o-quinone crossbond linkages. Tanning
probably occurs to some extent in all arthropods, but it is
particularly common in terrestrial forms such as insects
and arachnids.
The underlying epidermis or hypoepidermis is respon-
sible for the secretion of the cuticle. In addition to epithelial
cells, this tissue contains various unicellular glands, which
may have ducts to the surface of the cuticle. Beneath the
epidermis lies a distinct basement membrane, which forms
the outer boundary of the main body cavity or haemocoel.
Arthropod Physiology
Arthropod physiology is in large part dictated by two fea-
tures of the cuticle: first that it is relatively impermeable,
and second that it usually cannot stretch or contract. Since
Seta
Opening of
tegumental
gland duct
Cuticle
Membranous
layer
Tegumental
gland
3
 Arthropoda (Arthropods)
a muscular, flexible body wall cannot be used to enhance
blood flow, a muscular, pumping heart is necessary for
circulation. On the other hand, since there are no closed
coelomic spaces, an open circulatory system with a ha-
emocoel is possible. Respiratory pigments, particularly
haemocyanin, are common in larger forms. With regard to
excretion, nephridia with open nephrostomes are not fea-
sible – the blood of the haemocoel cannot be directly
drained to the outside! Excretory structures must be inter-
nally closed and are often coelomic remnants associated
with particular segments, although the identity of these
segments may vary between different taxa, e.g. excretory
structures are found on head segments in crustaceans and
leg segments in arachnids. These efficient mechanisms of
osmoregulation, coupled with the impermeable cuticle,
contribute to the success of arthropods in freshwater and
terrestrial environments. On the other hand, except per-
haps in tiny arthropods with high surface-to-volume ra-
tios, the largely impermeable cuticle does not allow gas
exchange through the body wall so specialized structures
have been developed. These structures take very different
forms in marine and aquatic versus terrestrial groups.
Larger marine and aquatic forms usually possess blood-
filled, thin-walled cuticular evaginations that serve as gills.
Terrestrial forms, on the other hand, have cuticular in-
vaginations of various types, which allow oxygen to enter
solution without excessive water loss.
The characteristic arthropod cuticle has its most dra-
matic effects on mechanisms of growth. A rigid exoskel-
eton precludes gradual changes in body size; consequently,
growth takes place episodically by means of the process of
moulting or ecdysis. Many features of this process may be
shared derived features of the ecdysozoa; certainly, be-
cause of the elaborate structure and function of the ar-
thropod cuticle, these moulting cycles dominate the
biology of arthropods. Actual tissue growth occurs dur-
ing the intermoult stages (instars). When the exoskeleton is
‘filled’ by this cell multiplication, the arthropod enters the
premoult stage (proecdysis). Certain epidermal glands se-
crete enzymes that digest the old endocuticle, and the ep-
idermis begins to separate from the exoskeleton. Beneath
the old exoskeleton the epidermis secretes a new, soft cu-
ticle. Subsequent to these events, actual ecdysis occurs. The
old cuticle splits, usually in a characteristic manner for
particular taxa, and the freshly moulted arthropod pulls
itself out. Immediately, this individual swells by the rapid
uptake of water or air. The postmoult stage (postecdysis)
commences with the hardening of the new exoskeleton by
mineralization, or sclerotization, or both. It should be ap-
parent that the many advantages of the exoskeleton are lost
until the new exoskeleton has hardened; for example, even
locomotion is difficult. Natural selection has no doubt
streamlined the physiology of arthropod moulting. Nev-
ertheless, as fossils in the Burgess Shale attest, the first
arthropods appear to have moulted much as modern ones
do. See also: Chelicerata (arachnids, including spiders,
4
mites and scorpions); Ecology of invertebrate nutrition;
Regeneration in crustaceans and insects
Since the moulting process involves the synchronization
of various organism-wide physiological processes, it is no
surprise that this process is hormonally controlled. Ecdy-
sone is responsible for the initiation of proecdysis. At least
in crustaceans, the secretion of this hormone is controlled
in turn by moult-inhibiting hormone (MIH), which is pro-
duced by neurosecretory cells during intermoult stages.
Sensory input from the nervous system mediates external
(e.g. tidal cycles) or internal (e.g. soft tissue growth) stimuli
and downregulates MIH, leading to the secretion of ecdy-
sone and the initiation of active moulting.
While compound eyes occur in some lophotrochozoan
groups (e.g. annelids and molluscs), these are probably in-
dependently derived from similar structures in arthropods.
Indeed, studies of ostracods suggest that even among ar-
thropods, compound eyes may have been independently
derived. Certainly, the physiology of these eyes is a major
topic in the study of arthropods. As their name implies,
compound eyes include several or many distinct photore-
ceptive units, the ommatidia. Each ommatidium has its
own field of vision, and each is supplied with its own nerve
fibres that lead to the optic nerve. The visual fields of the
ommatidia overlap to some degree. Thus the image that is
formed by a compound eye differs greatly from that pro-
duced by a complex camera-type eye such as that found in
cephalopods and vertebrates. In particular, compound
eyes are useful for detecting and analysing movement, be-
cause a shift in the position of an observed object causes
changes in the nervous impulses from several ommatidia.
See also: Sensory processing in invertebrate motor systems
The Impact of Arthropods in the
Modern World
Arthropods are inescapable in modern human life – as
food, as pests, as carriers of disease and as ‘model systems’
in modern biology.
Drosophila melanogaster, the fruitfly, is one of the prin-
cipal ‘model systems’ of modern biology. Much insight
into biology in general and human biomedicine in partic-
ular has been and will continue to be provided by D. me-
lanogaster. For instance, the homeobox, a sequence of 60
amino acids that forms a helix-turn-helix motif and binds
to deoxyribonucleic acid (DNA), was first discovered in
D. melanogaster. This sequence characterizes a group of
regulatory genes that has now been found to be widespread
in animals. These genes act as ‘master switches’ in a number
of developmental processes. Notably, D. melanogaster and
the nematode Caenorhabditis elegans represent the first
metazoa to have their entire genomes sequenced. The study
of these two creatures will no doubt continue to enhance
our understanding of genetic mechanisms; nevertheless,

the general applicability of these two model systems may be
limited by their close relationship as ecdysozoa. See also:
Caenorhabditis elegans as an experimental organism;
Regulatory genes in plant development: homeobox
Insects are carriers of many human diseases. Mosquitoes
in particular are vectors for malaria, yellow fever, viral
encephalitis, dengue and other serious illnesses. Malaria,
one of the most widespread human diseases, illustrates the
sometimes complex process of disease transmission. When
a female Anopheles mosquito feeds on human blood, she
releases an immature stage of the protist Plasmodium.
These protists multiply and mature within the human liver
and blood. Mosquitoes feeding on humans now pick up
mature Plasmodium, which then mate in the gut of the
mosquito. Immature forms are produced within the mos-
quito, thus completing the life cycle and leading to subse-
quent human infection. Now that the genomes of both the
protist parasite and the mosquito vector have been se-
quenced, a more complete understanding of the biology
of these organisms should lead to progress in stopping
malaria. See also: Malaria; Plasmodium
Roughly 10 000 years ago as human agriculture devel-
oped in several parts of the world, one of the goals of these
first plant breeders was probably to improve the produc-
tion and palatability of these first crops. Inadvertently, in
the process, the first conflicts between humans and insects
were begun, in that selection for improved production and
palatability also rendered these crops more vulnerable to
insect herbivores, or ‘pests’. This early human–insect con-
flict is manifest even in the Bible (e.g. the plague of locusts
in Exodus). In the modern world, billions of dollars are
spent annually to protect human crops. It is simply im-
possible for humans to grow food without protecting it
from insects either with extrinsic pesticide use (e.g. di-
chlorodiphenyltrichloroethane (DDT)) or with intrinsic
pesticides, or usually with both. Intrinsic pesticides, i.e.
those produced directly and ‘naturally’ by plants them-
selves, may pose serious health risks for humans. It is es-
timated that humans ingest roughly 10 000 different kinds
of these ‘natural’ pesticides and their breakdown products.
For instance, when cabbage is eaten, 49 natural pesticides
are ingested, including glucosinolates, indoles, is-
othiocyanates, cyanides, terpenes and phenols. These and
other compounds are widespread in domestic crops, and
function largely to repel insect herbivores. Presently, there
is considerable debate about the use of genetically modified
crops, which also intrinsically produce toxins to repel in-
sect herbivores (e.g. Bt corn). Environmental pollution re-
lated to pesticide use, human disease caused by natural and
synthetic dietary pesticides, and social conflicts over ge-
netically modified food all are ultimately due to the battle
between humans and insects over food. It is of note that
despite the best human efforts, insects consume roughly
one-third of human harvests or potential harvests annu-
ally. See also: Plant breeding and crop improvement; Plant
virus transmission by insects
Arthropoda (Arthropods)
On a more positive note, honeybees alone pollinate bil-
lions of dollars of crops annually, not to mention produc-
ing millions of dollars worth of honey. Also, since humans
do not directly farm the ocean for crops, and since crus-
taceans are largely marine, this group of arthropods is a
major source of food for humans. Human fisheries of
crustacean species abound, for instance, for crabs (e.g.
snow and tanner crabs), crab-like species (the Alaska king
crab and the South American centolla), shrimp and their
relatives, and lobster. In the case of the American lobster,
Homarus americanus, record catch amounts have occurred
in recent years. Although the reasons for this are obscure,
the overfishing of predatory fish species may be a contrib-
uting factor. Other fisheries have precipitously declined,
for example, that of the red Alaskan king crab (Paralith-
odes camtschatica). In some areas, ‘farming’ of freshwater
shrimp is occurring, sometimes with detrimental environ-
mental effects. Some human fisheries of nonarthropods are
indirectly based on crustaceans, for instance, those of krill-
eating whales, squid and fish in the ocean surrounding the
Antarctic. While clearly the bulk of human diet is derived
from land-based agriculture, direct and indirect fisheries of
crustaceans have an impact disproportionate to their size
because they provide high-quality protein. Given the 10%
rule of ecological efficiency, replacing these fisheries with
additional land-based agriculture would be essentially im-
possible. See also: Fisheries management
Fossil History
It is not merely in the present day that arthropods are the
most abundant and diverse multicellular organisms; rath-
er, the dominance of arthropods can be seen throughout
the history of animal life, beginning with well-preserved
fossil assemblages from the Cambrian. For instance, the
Burgess Shale preserves the fauna from an ancient marine
reef about 525 million years ago, near the dawn of abun-
dant large-scale animal life. As with other Cambrian as-
semblages, trilobites such as Olenoides are common.
Modern arthropods are also represented. Crustaceans ap-
pear in the form of Canadaspis, chelicerates as Sanctacaris,
and onychophorans as Aysheaia. Nevertheless, there are
also many arthropods, including the most abundant fossil
Marrella, which do not so clearly fit into any of these (or
other) groups of arthropods. In fact, one of the liveliest
recent scientific debates focuses on these diverse Burgess
Shale arthropods, their phylogenetic affinities, their mor-
phological disparity, and their general significance in terms
of organismal evolution. See also: Burgess Shale; Cam-
brian radiation; Fossils and fossilization
While trilobites, in part because of their well-mineralized
exoskeleton, dominate other Cambrian and Ordovician
marine fossil beds, by the later Palaeozoic trilobites are less
common, eventually becoming extinct in the end-Permian
mass extinction about 250 million years ago. Merostomates
5
 Arthropoda (Arthropods)
(xiphosurans and eurypterids) are known from Ordovician
marine deposits and flourished in the Silurian and Devon-
ian. Eurypterids, the water scorpions, were often quite large
and in some cases were common in freshwater and possibly
terrestrial environments. While the eurypterids became en-
tirely extinct in the Permian, a few species of xiphosurans
are still extant (Figure 2). See also: Extinction: end-Permian
mass extinction; Trilobita (trilobites)
Scorpions, perhaps a sister group of the eurypterids, are
among the most ancient terrestrial arthropods, invading
land prior to the Carboniferous. Insects also invaded ter-
restrial habitats at this time, although Devonian insect
fossils are rare. Winged insects are common as Carboni-
ferous fossils, suggesting that wings arose soon after insects
colonized land. Indeed, much of the success of insects is
attributable to their wings. Early insect fossils do not elu-
cidate the structure that wings evolved from, although
studies of developmental regulatory mechanisms in mod-
ern insects suggest that they may have evolved from the
basic limb structure. Generally, insect diversity coevolved
with terrestrial plants, particularly angiosperms. Perhaps
appropriately, amber is a major source of exquisitely pre-
served fossil insects. While there has been considerable
popular and scientific interest in obtaining ancient DNA
from insects in amber, initial reports were overly optimistic
concerning the likelihood of success in this regard.
As with insects and their wings, the number, morphol-
ogy and function of body segments and limbs have re-
flected the diversification of the arthropods throughout the
Phanerozoic. With marine forms this was particularly
apparent during the Mesozoic. At this time, marine crus-
taceans and their prey radiated extensively. Various crus-
tacean groups (e.g. stomatopods and decapods) developed
characteristic appendages that allowed them to become
much more sophisticated in subduing their prey. In turn,
groups of prey such as molluscs developed better anti-
predator defences, e.g. in features of their shells such as
thickness and sculpture. See also: Mollusca (molluscs)
Phylogenetic Controversy – One Group
or Many?
Discussions of the evolution of arthropods have been tied
to the premise that annelids and panarthropods (Arthro-
poda + Tardigrada + Onychophora) are sister taxa with-
in the larger ‘Articulata’ taxon. A plausible stem lineage for
these sister taxa would be a segmented, soft-bodied, worm-
like creature with serially arranged coelomic spaces and
well-developed circular and longitudinal muscles. If such a
creature derived the major arthropod feature – a hard,
jointed exoskeleton – a number of structural and func-
tional correlates would have to evolve simultaneously.
Circular and longitudinal muscles would no longer be use-
ful in locomotion and would be lost, while striated muscles
6
organized as intersegmental bands would evolve (Figure 1).
The coelom could no longer be used as a hydrostatic skel-
eton and would be lost, while an open circulatory system
and a haemocoel would evolve. The latter would require a
muscular heart and a closed excretory system. Moulting or
ecdysis of the rigid exoskeleton would be required for
growth. Adaptations for coping with osmotic and ionic
stress, and for gas exchange, would evolve as appropriate.
The evolution of these features collectively has been termed
‘arthropodization’, and the controversy over arthropod
monophyly has focused on the very real possibility that this
evolutionary scenario may have occurred more than once,
i.e. that this suite of functionally interconnected traits
might be evolved independently in different groups of ar-
thropods. See also: Annelida (segment worms)
This controversy has a long history, and numerous spe-
cific hypotheses of arthropod polyphyly have been pro-
posed. For instance, some have argued for a Myriapoda
+Hexapoda +Onychophora clade (‘Uniramia’) with
another clade comprising the trilobites, crustaceans and
chelicerates. Other phylogenetic hypotheses describe a
paraphyletic Arthropoda. See also: Myriapoda (including
centipedes and millipedes)
Nevertheless, the controversy over arthropod systema-
tics has been completely recast by the recent ‘ecdysozoan
hypothesis’, i.e. panarthropods and annelids are not sister
taxa, but belong to distinct clades of metazoa, presently
termed Ecdysozoa and Lophotrochozoa. Within the
Ecdysozoa, the sister group of the panarthropods now ap-
pears to be the clade containing the nematodes and the
nematomorphs. By this hypothesis, evolutionary scenarios
deriving arthropods from annelid-like ancestors via the
process of ‘arthropodization’ would be untenable. The
ecdysozoan hypothesis nevertheless remains controversial
(see the next section). See also: Echiura (unsegmented
protostome worms); Nematoda (roundworms)
Modern Phylogeny
In what some have characterized as a debate between
‘molecules and morphology’, nucleotide sequence data,
particularly of 18S ribosomal DNA (rDNA), suggest that
the relationship between annelids and panarthropods is
unlikely to be close. Rather, these molecular data indicate
that arthropods and allied taxa form the Ecdysozoa, a
clade including other moulting animals. In particular, the
Ecdysozoa emerges when rapidly evolving nematode se-
quences (which can lead to long-branch attraction and an
apparent basal position for the nematodes) are removed
from the phylogenetic analysis. Nevertheless, some careful
morphological analyses have also raised questions about
the close relationship between panarthropods and anne-
lids, while at the same time sequence data from a number
of genes neither supports nor contradicts the ecdysozoan
 Arthropoda (Arthropods)

Table 1 A classification of the phylum Arthropoda and their evidence (e.g. that of development) can contribute to this
close relatives synthesis.
Panarthropoda
Tardigrada Further Reading
Onychophora Aguinaldo AMA, Turbeville JM, Linford LS et al. (1997) Evidence for a
Arthropoda clade of nematodes, arthropods and other moulting animals. Nature
Trilobita 387: 489–493.
Chelicerata Berenbaum MR (1995) Bugs in the System: Insects and their Impact on
Merostomata Human Affairs. New York: Addison-Wesley.
Arachnida Brusca RC and Brusca GJ (2003) Invertebrates, 2nd edn, Sunderland,
Pycnogonida MA: Sinauer.
Copley RR, Aloy P, Russel RB and Telford MJ (2004) Systematic
Mandibulata searches for molecular synapomorphies in model metazoan genomes
Myriapoda give some support for Ecdysozoa after accounting for the idiosyncra-
Hexapoda sies of Caenorhabditis elegans. Evolution and Development 6: 164–169.
Crustacea Edgecomb GD (ed.) (1998) Arthropod Fossils and Phylogeny. New York:
Columbia University Press.
Fortey RA and Thomas RH (eds) (1998) Arthropod Relationships. Sys-
tematics Association Special Volume Series 55. London: Chapman
and Hall.
hypothesis. In terms of relationships within the arthro- Lawrence PA (1992) The Makings of a Fly. Oxford: Blackwell.
pods, morphological analyses have never reached a general McHugh D and Halanych KM (eds) (1998) Evolutionary relationships
consensus, and again molecular data have been and will of metazoan phyla: advances, problems, and approaches. American
Zoologist 38: 813–981.
continue to be crucial. The classification shown here
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