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Contents
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(Top)

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Etymology


Description

 
 Diversity

Segmentation

Exoskeleton

Moulting

Internal organs

Respiration and circulation

Nervous system

Excretory system


 Senses

Optical

Olfaction


Reproduction and development


Evolutionary history

 

Last common ancestor

Fossil record

Evolutionary relationships to other animal phyla

Evolution of fossil arthropods

 

Evolution and classification of living arthropods


Interaction with humans


See also


Notes

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References

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o

Bibliography

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Arthropod

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From Wikipedia, the free encyclopedia

Not to be confused with Anthropod.
Arthropoda
Temporal range: 538.8 –0 Ma
PreꞒ
Ꞓ
O
S
D
C
P
T
J
K
Pg
N
Earliest Cambrian (Fortunian)–Recent
 Scientific classification
Eukary
Domain:
ota
Anima
Kingdom:
lia
Eumet
Subkingdom:
azoa
ParaH
Clade:
oxozoa
Bilater
Clade:
ia
Nephr
Clade:
ozoa
Protost
(unranked):
omia
Ecdyso
Superphylum:
zoa
Panart
(unranked): hropod
a
Tactop
(unranked):
oda
Arthro
poda
Phylum: Gravenh
orst,
1843[1][2]
Subphyla, unplaced genera,
and classes
 Diversity
around 1,170,000 species.
Synonyms
Condylipoda Latreille, 1802

Arthropods (/ˈɑːrθrəpɒd/) are invertebrates in the phylum Arthropoda. They possess an

exoskeleton with a cuticle made of chitin, often mineralised with calcium carbonate, a body with
differentiated (metameric) segments, and paired jointed appendages. In order to keep growing,
they must go through stages of moulting, a process by which they shed their exoskeleton to
reveal a new one. They are an extremely diverse group, with up to 10 million species.

Haemolymph is the analogue of blood for most arthropods. An arthropod has an open circulatory
system, with a body cavity called a haemocoel through which haemolymph circulates to the
interior organs. Like their exteriors, the internal organs of arthropods are generally built of
repeated segments. Their nervous system is "ladder-like", with paired ventral nerve cords
running through all segments and forming paired ganglia in each segment. Their heads are
formed by fusion of varying numbers of segments, and their brains are formed by fusion of the
ganglia of these segments and encircle the esophagus. The respiratory and excretory systems of
arthropods vary, depending as much on their environment as on the subphylum to which they
belong.

Arthropods use combinations of compound eyes and pigment-pit ocelli for vision. In most
species, the ocelli can only detect the direction from which light is coming, and the compound
eyes are the main source of information, but the main eyes of spiders are ocelli that can form
images and, in a few cases, can swivel to track prey. Arthropods also have a wide range of
chemical and mechanical sensors, mostly based on modifications of the many bristles known as
setae that project through their cuticles. Similarly, their reproduction and development are
varied; all terrestrial species use internal fertilization, but this is sometimes by indirect transfer of
the sperm via an appendage or the ground, rather than by direct injection. Aquatic species use
either internal or external fertilization. Almost all arthropods lay eggs, with many species giving
birth to live young after the eggs have hatched inside the mother; but a few are genuinely
viviparous, such as aphids. Arthropod hatchlings vary from miniature adults to grubs and
caterpillars that lack jointed limbs and eventually undergo a total metamorphosis to produce the
adult form. The level of maternal care for hatchlings varies from nonexistent to the prolonged
care provided by social insects.

The evolutionary ancestry of arthropods dates back to the Cambrian period. The group is
generally regarded as monophyletic, and many analyses support the placement of arthropods
with cycloneuralians (or their constituent clades) in a superphylum Ecdysozoa. Overall, however,
the basal relationships of animals are not yet well resolved. Likewise, the relationships between
various arthropod groups are still actively debated. Today, arthropods contribute to the human
food supply both directly as food, and more importantly, indirectly as pollinators of crops. Some
species are known to spread severe disease to humans, livestock, and crops.

Etymology
The word arthropod comes from the Greek ἄρθρον árthron 'joint', and πούς pous (gen. ποδός
podos) 'foot' or 'leg', which together mean "jointed leg",[19] with the word "arthropodes" initially
used in anatomical descriptions by Barthélemy Charles Joseph Dumortier published in 1832.[1]
The designation "Arthropoda" appears to have been first used in 1843 by the German zoologist
Johann Ludwig Christian Gravenhorst (1777–1857).[20][1] The origin of the name has been the
subject of considerable confusion, with credit often given erroneously to Pierre André Latreille
or Karl Theodor Ernst von Siebold instead, among various others.[1]

Terrestrial arthropods are often called bugs.[Note 1] The term is also occasionally extended to
colloquial names for freshwater or marine crustaceans (e.g., Balmain bug, Moreton Bay bug,
mudbug) and used by physicians and bacteriologists for disease-causing germs (e.g., superbugs),
[23]
but entomologists reserve this term for a narrow category of "true bugs", insects of the order
Hemiptera.[23]

Description
Arthropods are invertebrates with segmented bodies and jointed limbs.[24] The exoskeleton or
cuticles consists of chitin, a polymer of N-Acetylglucosamine.[25] The cuticle of many
crustaceans, beetle mites, the clades Penetini and Archaeoglenini inside the beetle subfamily
Phrenapatinae,[26] and millipedes (except for bristly millipedes) is also biomineralized with
calcium carbonate. Calcification of the endosternite, an internal structure used for muscle
attachments, also occur in some opiliones,[27] and the pupal cuticle of the fly Bactrocera dorsalis
contains calcium phosphate.[28]

Diversity

Protaetia cuprea (copper chafer). Beetles are the largest and

most diverse order of arthropods.
Arthropoda is the largest animal phylum with the estimates of the number of arthropod species
varying from 1,170,000 to 5 to 10 million and accounting for over 80 per cent of all known
living animal species.[29][30] One arthropod sub-group, the insects, includes more described
species than any other taxonomic class.[31] The total number of species remains difficult to
determine. This is due to the census modeling assumptions projected onto other regions in order
to scale up from counts at specific locations applied to the whole world. A study in 1992
estimated that there were 500,000 species of animals and plants in Costa Rica alone, of which
365,000 were arthropods.[31]

They are important members of marine, freshwater, land and air ecosystems and one of only two
major animal groups that have adapted to life in dry environments; the other is amniotes, whose
living members are reptiles, birds and mammals.[32] Both the smallest and largest arthropods are
crustaceans. The smallest belong to the class Tantulocarida, some of which are less than 100
micrometres (0.0039 in) long.[33] The largest are species in the class Malacostraca, with the legs
of the Japanese spider crab potentially spanning up to 4 metres (13 ft)[34] and the American
lobster reaching weights over 20 kg (44 lbs).

Segmentation

Head
_______________________
Thorax
_______________________
Abdomen
_______________________

Segments and tagmata of an arthropod[32]

 Structure of a biramous appendage.[35]

The embryos of all arthropods are segmented, built from a series of repeated modules. The last
common ancestor of living arthropods probably consisted of a series of undifferentiated
segments, each with a pair of appendages that functioned as limbs. However, all known living
and fossil arthropods have grouped segments into tagmata in which segments and their limbs are
specialized in various ways.[32]

The three-part appearance of many insect bodies and the two-part appearance of spiders is a
result of this grouping.[36] There are no external signs of segmentation in mites.[32] Arthropods
also have two body elements that are not part of this serially repeated pattern of segments, an
ocular somite at the front, where the mouth and eyes originated,[32][37] and a telson at the rear,
behind the anus.

Originally it seems that each appendage-bearing segment had two separate pairs of appendages:
an upper, unsegmented exite and a lower, segmented endopod. These would later fuse into a
single pair of biramous appendages united by a basal segment (protopod or basipod), with the
upper branch acting as a gill while the lower branch was used for locomotion.[38][39][35] The
appendages of most crustaceans and some extinct taxa such as trilobites have another segmented
branch known as exopods, but whether these structures have a single origin remain controversial.
[40][41][35]
In some segments of all known arthropods the appendages have been modified, for
example to form gills, mouth-parts, antennae for collecting information,[36] or claws for grasping;
[42]
arthropods are "like Swiss Army knives, each equipped with a unique set of specialized
tools."[32] In many arthropods, appendages have vanished from some regions of the body; it is
particularly common for abdominal appendages to have disappeared or be highly modified.[32]
Alignment of anterior body segments and appendages across various arthropod taxa, based on
the observations until the mid 2010s. Head regions in black.[37][43]

The most conspicuous specialization of segments is in the head. The four major groups of
arthropods – Chelicerata (sea spiders, horseshoe crabs and arachnids), Myriapoda (symphylan,
pauropods, millipedes and centipedes), Pancrustacea (oligostracans, copepods, malacostracans,
branchiopods, hexapods, etc.), and the extinct Trilobita – have heads formed of various
combinations of segments, with appendages that are missing or specialized in different ways. [32]
Despite myriapods and hexapods both having similar head combinations, hexapods are deeply
nested within crustacea while myriapods are not, so these traits are believed to have evolved
separately. In addition, some extinct arthropods, such as Marrella, belong to none of these
groups, as their heads are formed by their own particular combinations of segments and
specialized appendages.[44]

Working out the evolutionary stages by which all these different combinations could have
appeared is so difficult that it has long been known as "The arthropod head problem".[45] In 1960,
R. E. Snodgrass even hoped it would not be solved, as he found trying to work out solutions to
be fun.[Note 2]

Exoskeleton

Main article: Arthropod exoskeleton

 Illustration of an idealized arthropod exoskeleton.

Arthropod exoskeletons are made of cuticle, a non-cellular material secreted by the epidermis.[32]
Their cuticles vary in the details of their structure, but generally consist of three main layers: the
epicuticle, a thin outer waxy coat that moisture-proofs the other layers and gives them some
protection; the exocuticle, which consists of chitin and chemically hardened proteins; and the
endocuticle, which consists of chitin and unhardened proteins. The exocuticle and endocuticle
together are known as the procuticle.[47] Each body segment and limb section is encased in
hardened cuticle. The joints between body segments and between limb sections are covered by
flexible cuticle.[32]

The exoskeletons of most aquatic crustaceans are biomineralized with calcium carbonate
extracted from the water. Some terrestrial crustaceans have developed means of storing the
mineral, since on land they cannot rely on a steady supply of dissolved calcium carbonate.[48]
Biomineralization generally affects the exocuticle and the outer part of the endocuticle. [47] Two
recent hypotheses about the evolution of biomineralization in arthropods and other groups of
animals propose that it provides tougher defensive armor,[49] and that it allows animals to grow
larger and stronger by providing more rigid skeletons;[50] and in either case a mineral-organic
composite exoskeleton is cheaper to build than an all-organic one of comparable strength.[50][51]

The cuticle may have setae (bristles) growing from special cells in the epidermis. Setae are as
varied in form and function as appendages. For example, they are often used as sensors to detect
air or water currents, or contact with objects; aquatic arthropods use feather-like setae to increase
the surface area of swimming appendages and to filter food particles out of water; aquatic
insects, which are air-breathers, use thick felt-like coats of setae to trap air, extending the time
they can spend under water; heavy, rigid setae serve as defensive spines.[32]

Although all arthropods use muscles attached to the inside of the exoskeleton to flex their limbs,
some still use hydraulic pressure to extend them, a system inherited from their pre-arthropod
ancestors;[52] for example, all spiders extend their legs hydraulically and can generate pressures
up to eight times their resting level.[53]

Moulting

Main article: Ecdysis

Cicada climbing out of its exuviae while attached to tree

The exoskeleton cannot stretch and thus restricts growth. Arthropods, therefore, replace their
exoskeletons by undergoing ecdysis (moulting), or shedding the old exoskeleton, the exuviae,
after growing a new one that is not yet hardened. Moulting cycles run nearly continuously until
an arthropod reaches full size. The developmental stages between each moult (ecdysis) until
sexual maturity is reached is called an instar. Differences between instars can often be seen in
altered body proportions, colors, patterns, changes in the number of body segments or head
width. After moulting, i.e. shedding their exoskeleton, the juvenile arthropods continue in their
life cycle until they either pupate or moult again.[54]

In the initial phase of moulting, the animal stops feeding and its epidermis releases moulting
fluid, a mixture of enzymes that digests the endocuticle and thus detaches the old cuticle. This
phase begins when the epidermis has secreted a new epicuticle to protect it from the enzymes,
and the epidermis secretes the new exocuticle while the old cuticle is detaching. When this stage
is complete, the animal makes its body swell by taking in a large quantity of water or air, and this
makes the old cuticle split along predefined weaknesses where the old exocuticle was thinnest. It
commonly takes several minutes for the animal to struggle out of the old cuticle. At this point,
the new one is wrinkled and so soft that the animal cannot support itself and finds it very difficult
to move, and the new endocuticle has not yet formed. The animal continues to pump itself up to
stretch the new cuticle as much as possible, then hardens the new exocuticle and eliminates the
excess air or water. By the end of this phase, the new endocuticle has formed. Many arthropods
then eat the discarded cuticle to reclaim its materials.[54]

Because arthropods are unprotected and nearly immobilized until the new cuticle has hardened,
they are in danger both of being trapped in the old cuticle and of being attacked by predators.
Moulting may be responsible for 80 to 90% of all arthropod deaths.[54]

Internal organs

= heart
= gut
= brain / ganglia
O = eye

Basic arthropod body structure

Arthropod bodies are also segmented internally, and the nervous, muscular, circulatory, and
excretory systems have repeated components.[32] Arthropods come from a lineage of animals that
have a coelom, a membrane-lined cavity between the gut and the body wall that accommodates
the internal organs. The strong, segmented limbs of arthropods eliminate the need for one of the
coelom's main ancestral functions, as a hydrostatic skeleton, which muscles compress in order to
change the animal's shape and thus enable it to move. Hence the coelom of the arthropod is
reduced to small areas around the reproductive and excretory systems. Its place is largely taken
by a hemocoel, a cavity that runs most of the length of the body and through which blood flows.
[55]

Respiration and circulation

See also: Hemolymph and hemocyte

 Respiration and circulation in a myodocopid ostracod.
Simplified transverse section through anterior body and carapace, showing gaseous diffusion
through the inner lamella of the carapace (yellow arrows)

Arthropods have open circulatory systems. Most have a few short, open-ended arteries. In
chelicerates and crustaceans, the blood carries oxygen to the tissues, while hexapods use a
separate system of tracheae. Many crustaceans and a few chelicerates and tracheates use
respiratory pigments to assist oxygen transport. The most common respiratory pigment in
arthropods is copper-based hemocyanin; this is used by many crustaceans and a few centipedes.
A few crustaceans and insects use iron-based hemoglobin, the respiratory pigment used by
vertebrates. As with other invertebrates, the respiratory pigments of those arthropods that have
them are generally dissolved in the blood and rarely enclosed in corpuscles as they are in
vertebrates.[55]

The heart is a muscular tube that runs just under the back and for most of the length of the
hemocoel. It contracts in ripples that run from rear to front, pushing blood forwards. Sections not
being squeezed by the heart muscle are expanded either by elastic ligaments or by small muscles,
in either case connecting the heart to the body wall. Along the heart run a series of paired ostia,
non-return valves that allow blood to enter the heart but prevent it from leaving before it reaches
the front.[55]

Arthropods have a wide variety of respiratory systems. Small species often do not have any,
since their high ratio of surface area to volume enables simple diffusion through the body surface
to supply enough oxygen. Crustacea usually have gills that are modified appendages. Many
arachnids have book lungs.[56] Tracheae, systems of branching tunnels that run from the openings
in the body walls, deliver oxygen directly to individual cells in many insects, myriapods and
arachnids.[57]

Nervous system
 Central nervous system of a nectiopod remipede, showing the
presence of both deutocerebrum (dc) and ventral nerve cord (vnc) organized by segmented
ganglia.

Living arthropods have paired main nerve cords running along their bodies below the gut, and in
each segment the cords form a pair of ganglia from which sensory and motor nerves run to other
parts of the segment. Although the pairs of ganglia in each segment often appear physically
fused, they are connected by commissures (relatively large bundles of nerves), which give
arthropod nervous systems a characteristic "ladder-like" appearance. The brain is in the head,
encircling and mainly above the esophagus. It consists of the fused ganglia of the acron and one
or two of the foremost segments that form the head – a total of three pairs of ganglia in most
arthropods, but only two in chelicerates, which do not have antennae or the ganglion connected
to them. The ganglia of other head segments are often close to the brain and function as part of it.
In insects these other head ganglia combine into a pair of subesophageal ganglia, under and
behind the esophagus. Spiders take this process a step further, as all the segmental ganglia are
incorporated into the subesophageal ganglia, which occupy most of the space in the
cephalothorax (front "super-segment").[58]

Excretory system

There are two different types of arthropod excretory systems. In aquatic arthropods, the end-
product of biochemical reactions that metabolise nitrogen is ammonia, which is so toxic that it
needs to be diluted as much as possible with water. The ammonia is then eliminated via any
permeable membrane, mainly through the gills.[56] All crustaceans use this system, and its high
consumption of water may be responsible for the relative lack of success of crustaceans as land
animals.[59] Various groups of terrestrial arthropods have independently developed a different
system: the end-product of nitrogen metabolism is uric acid, which can be excreted as dry
material; the Malpighian tubule system filters the uric acid and other nitrogenous waste out of
the blood in the hemocoel, and dumps these materials into the hindgut, from which they are
expelled as feces.[59] Most aquatic arthropods and some terrestrial ones also have organs called
nephridia ("little kidneys"), which extract other wastes for excretion as urine.[59]

Senses
 Long bristles (setae) of a Tliltocatl albopilosus tarantula

The stiff cuticles of arthropods would block out information about the outside world, except that
they are penetrated by many sensors or connections from sensors to the nervous system. In fact,
arthropods have modified their cuticles into elaborate arrays of sensors. Various touch sensors,
mostly setae, respond to different levels of force, from strong contact to very weak air currents.
Chemical sensors provide equivalents of taste and smell, often by means of setae. Pressure
sensors often take the form of membranes that function as eardrums, but are connected directly
to nerves rather than to auditory ossicles. The antennae of most hexapods include sensor
packages that monitor humidity, moisture and temperature.[60]

Most arthropods lack balance and acceleration sensors, and rely on their eyes to tell them which
way is up. The self-righting behavior of cockroaches is triggered when pressure sensors on the
underside of the feet report no pressure. However, many malacostracan crustaceans have
statocysts, which provide the same sort of information as the balance and motion sensors of the
vertebrate inner ear.[60]

The proprioceptors of arthropods, sensors that report the force exerted by muscles and the degree
of bending in the body and joints, are well understood. However, little is known about what other
internal sensors arthropods may have.[60]

Optical

Main article: Arthropod eye

 Arthropod eyes

Head of a wasp with three ocelli (center), and compound

eyes at the left and right

Most arthropods have sophisticated visual systems that include one or more usually both of
compound eyes and pigment-cup ocelli ("little eyes"). In most cases ocelli are only capable of
detecting the direction from which light is coming, using the shadow cast by the walls of the cup.
However, the main eyes of spiders are pigment-cup ocelli that are capable of forming images,[60]
and those of jumping spiders can rotate to track prey.[61]

Compound eyes consist of fifteen to several thousand independent ommatidia, columns that are
usually hexagonal in cross section. Each ommatidium is an independent sensor, with its own
light-sensitive cells and often with its own lens and cornea.[60] Compound eyes have a wide field
of view, and can detect fast movement and, in some cases, the polarization of light.[62] On the
other hand, the relatively large size of ommatidia makes the images rather coarse, and compound
eyes are shorter-sighted than those of birds and mammals – although this is not a severe
disadvantage, as objects and events within 20 cm (8 in) are most important to most arthropods.[60]
Several arthropods have color vision, and that of some insects has been studied in detail; for
example, the ommatidia of bees contain receptors for both green and ultra-violet.[60]

Olfaction

Further information: Insect olfaction

Reproduction and development

Aphid giving birth to live young from an unfertilized egg

Harvestmen mating

A few arthropods, such as barnacles, are hermaphroditic, that is, each can have the organs of
both sexes. However, individuals of most species remain of one sex their entire lives.[63] A few
species of insects and crustaceans can reproduce by parthenogenesis, especially if conditions
favor a "population explosion". However, most arthropods rely on sexual reproduction, and
parthenogenetic species often revert to sexual reproduction when conditions become less
favorable.[64] The ability to undergo meiosis is widespread among arthropods including both
those that reproduce sexually and those that reproduce parthenogenetically.[65] Although meiosis
is a major characteristic of arthropods, understanding of its fundamental adaptive benefit has
long been regarded as an unresolved problem,[66] that appears to have remained unsettled.

Aquatic arthropods may breed by external fertilization, as for example horseshoe crabs do,[67] or
by internal fertilization, where the ova remain in the female's body and the sperm must somehow
be inserted. All known terrestrial arthropods use internal fertilization. Opiliones (harvestmen),
millipedes, and some crustaceans use modified appendages such as gonopods or penises to
transfer the sperm directly to the female. However, most male terrestrial arthropods produce
spermatophores, waterproof packets of sperm, which the females take into their bodies. A few
such species rely on females to find spermatophores that have already been deposited on the
ground, but in most cases males only deposit spermatophores when complex courtship rituals
look likely to be successful.[63]
 The nauplius larva of a penaeid shrimp

Most arthropods lay eggs,[63] but scorpions are ovoviviparous: they produce live young after the
eggs have hatched inside the mother, and are noted for prolonged maternal care.[68] Newly born
arthropods have diverse forms, and insects alone cover the range of extremes. Some hatch as
apparently miniature adults (direct development), and in some cases, such as silverfish, the
hatchlings do not feed and may be helpless until after their first moult. Many insects hatch as
grubs or caterpillars, which do not have segmented limbs or hardened cuticles, and
metamorphose into adult forms by entering an inactive phase in which the larval tissues are
broken down and re-used to build the adult body.[69] Dragonfly larvae have the typical cuticles
and jointed limbs of arthropods but are flightless water-breathers with extendable jaws. [70]
Crustaceans commonly hatch as tiny nauplius larvae that have only three segments and pairs of
appendages.[63]

Evolutionary history
See also: Phylogeny of insects

Last common ancestor

Based on the distribution of shared plesiomorphic features in extant and fossil taxa, the last
common ancestor of all arthropods is inferred to have been as a modular organism with each
module covered by its own sclerite (armor plate) and bearing a pair of biramous limbs.[71]
However, whether the ancestral limb was uniramous or biramous is far from a settled debate.
This Ur-arthropod had a ventral mouth, pre-oral antennae and dorsal eyes at the front of the
body. It was assumed to have been a non-discriminatory sediment feeder, processing whatever
sediment came its way for food,[71] but fossil findings hint that the last common ancestor of both
arthropods and priapulida shared the same specialized mouth apparatus; a circular mouth with
rings of teeth used for capturing animal prey.[72]

Fossil record
 Marrella, one of the puzzling arthropods from the Burgess
Shale

It has been proposed that the Ediacaran animals Parvancorina and Spriggina, from around
555 million years ago, were arthropods,[73][74][75] but later study shows that their affinities of being
origin of arthropods are not reliable.[76] Small arthropods with bivalve-like shells have been
found in Early Cambrian fossil beds dating 541 to 539 million years ago in China and Australia.
[77][78][79][80]
The earliest Cambrian trilobite fossils are about 520 million years old, but the class
was already quite diverse and worldwide, suggesting that they had been around for quite some
time.[81] In the Maotianshan shales, which date back to 518 million years ago, arthropods such as
Kylinxia and Erratus have been found that seem to represent transitional fossils between stem
(e.g. Radiodonta such as Anomalocaris) and true arthropods.[82][3][39] Re-examination in the 1970s
of the Burgess Shale fossils from about 505 million years ago identified many arthropods, some
of which could not be assigned to any of the well-known groups, and thus intensified the debate
about the Cambrian explosion.[83][84][85] A fossil of Marrella from the Burgess Shale has provided
the earliest clear evidence of moulting.[86]

Kylinxia may be a key transitional fossil between stem-

arthropods and true arthropods.[82] Yicaris is one of the earliest

crustaceans that have been discovered.

The earliest fossil of likely pancrustacean larvae date from about 514 million years ago in the
Cambrian, followed by unique taxa like Yicaris and Wujicaris.[87] The purported
pancrustacean/crustacean affinity of some cambrian arthropods (e.g. Phosphatocopina,
Bradoriida and Hymenocarine taxa like waptiids)[88][89][90] were disputed by subsequent studies, as
they might branch before the mandibulate crown-group.[87] Within the pancrustacean crown-
group, only Malacostraca, Branchiopoda and Pentastomida have Cambrian fossil records.[87]
Crustacean fossils are common from the Ordovician period onwards.[91] They have remained
almost entirely aquatic, possibly because they never developed excretory systems that conserve
water.[59]

Arthropods provide the earliest identifiable fossils of land animals, from about 419 million years
ago in the Late Silurian,[56] and terrestrial tracks from about 450 million years ago appear to have
been made by arthropods.[92] Arthropods possessed attributes that were easy coopted for life on
land; their existing jointed exoskeletons provided protection against desiccation, support against
gravity and a means of locomotion that was not dependent on water.[93] Around the same time the
aquatic, scorpion-like eurypterids became the largest ever arthropods, some as long as 2.5 m (8 ft
2 in).[94]

The oldest known arachnid is the trigonotarbid Palaeotarbus jerami, from about 420 million
years ago in the Silurian period.[95][Note 3] Attercopus fimbriunguis, from 386 million years ago in
the Devonian period, bears the earliest known silk-producing spigots, but its lack of spinnerets
means it was not one of the true spiders,[97] which first appear in the Late Carboniferous over
299 million years ago.[98] The Jurassic and Cretaceous periods provide a large number of fossil
spiders, including representatives of many modern families.[99] The oldest known scorpion is
Dolichophonus, dated back to 436 million years ago.[100] Lots of Silurian and Devonian scorpions
were previously thought to be gill-breathing, hence the idea that scorpions were primitively
aquatic and evolved air-breathing book lungs later on.[101] However subsequent studies reveal
most of them lacking reliable evidence for an aquatic lifestyle,[102] while exceptional aquatic taxa
(e.g. Waeringoscorpio) most likely derived from terrestrial scorpion ancestors.[103]

The oldest fossil record of hexapod is obscure, as most of the candidates are poorly preserved
and their hexapod affinities had been disputed. An iconic example is the Devonian Rhyniognatha
hirsti, dated at 396 to 407 million years ago, its mandibles are thought to be a type found only in
winged insects, which suggests that the earliest insects appeared in the Silurian period.[104]
However later study shows that Rhyniognatha most likely represent a myriapod, not even a
hexapod.[105] The unequivocal oldest known hexapod and insect is the springtail Rhyniella, from
about 410 million years ago in the Devonian period, and the palaeodictyopteran Delitzschala
bitterfeldensis, from about 325 million years ago in the Carboniferous period, respectively.[105]
The Mazon Creek lagerstätten from the Late Carboniferous, about 300 million years ago, include
about 200 species, some gigantic by modern standards, and indicate that insects had occupied
their main modern ecological niches as herbivores, detritivores and insectivores. Social termites
and ants first appear in the Early Cretaceous, and advanced social bees have been found in Late
Cretaceous rocks but did not become abundant until the Middle Cenozoic.[106]

Evolutionary relationships to other animal phyla

 The velvet worm (Onychophora) is closely related to
arthropods[107]

From 1952 to 1977, zoologist Sidnie Manton and others argued that arthropods are polyphyletic,
in other words, that they do not share a common ancestor that was itself an arthropod. Instead,
they proposed that three separate groups of "arthropods" evolved separately from common
worm-like ancestors: the chelicerates, including spiders and scorpions; the crustaceans; and the
uniramia, consisting of onychophorans, myriapods and hexapods. These arguments usually
bypassed trilobites, as the evolutionary relationships of this class were unclear. Proponents of
polyphyly argued the following: that the similarities between these groups are the results of
convergent evolution, as natural consequences of having rigid, segmented exoskeletons; that the
three groups use different chemical means of hardening the cuticle; that there were significant
differences in the construction of their compound eyes; that it is hard to see how such different
configurations of segments and appendages in the head could have evolved from the same
ancestor; and that crustaceans have biramous limbs with separate gill and leg branches, while the
other two groups have uniramous limbs in which the single branch serves as a leg.[108]

onychophorans
includes Aysheaia and Peripatus

armored lobopods
includes Hallucigenia and Microdictyon

dinocarids (s.l.) anomalocarid- includes modern tardigrades as

well as extinct animals like
like taxa (s.l.) Kerygmachela and Opabinia
anomalocarids (s.s.)
Anomalocari
s
arthropods includes
living groups
and
extinct forms
such as
trilobites
Simplified summary of Budd's (1996) "broad-scale" cladogram[107]
Further analysis and discoveries in the 1990s reversed this view, and led to acceptance that
arthropods are monophyletic, in other words they are inferred to share a common ancestor that
was itself an arthropod.[109][110] For example, Graham Budd's analyses of Kerygmachela in 1993
and of Opabinia in 1996 convinced him that these animals were similar to onychophorans and to
various Early Cambrian "lobopods", and he presented an "evolutionary family tree" that showed
these as "aunts" and "cousins" of all arthropods.[107][111] These changes made the scope of the term
"arthropod" unclear, and Claus Nielsen proposed that the wider group should be labelled
"Panarthropoda" ("all the arthropods") while the animals with jointed limbs and hardened
cuticles should be called "Euarthropoda" ("true arthropods").[112]

A contrary view was presented in 2003, when Jan Bergström and Hou Xian-guang argued that, if
arthropods were a "sister-group" to any of the anomalocarids, they must have lost and then re-
evolved features that were well-developed in the anomalocarids. The earliest known arthropods
ate mud in order to extract food particles from it, and possessed variable numbers of segments
with unspecialized appendages that functioned as both gills and legs. Anomalocarids were, by
the standards of the time, huge and sophisticated predators with specialized mouths and grasping
appendages, fixed numbers of segments some of which were specialized, tail fins, and gills that
were very different from those of arthropods. In 2006, they suggested that arthropods were more
closely related to lobopods and tardigrades than to anomalocarids.[113] In 2014, it was found that
tardigrades were more closely related to arthropods than velvet worms.[114]

Protostom
es
Chaetognatha

Spiralia (annelids, molluscs, brachiopods, etc.)

Ecdysoz
oa
Nematoida (nematodes and close relatives)

Scalidophora (priapulids and Kinorhyncha, and Loricifera)

Panarthrop
oda Onychophorans
Tactopo
da Tardigrades
Euarthropo Chelicerates
da
Relationships of Ecdysozoa to each other and to annelids, etc.,[115][failed verification] including
euthycarcinoids[116]

Higher up the "family tree", the Annelida have traditionally been considered the closest relatives
of the Panarthropoda, since both groups have segmented bodies, and the combination of these
groups was labelled Articulata. There had been competing proposals that arthropods were closely
related to other groups such as nematodes, priapulids and tardigrades, but these remained
minority views because it was difficult to specify in detail the relationships between these
groups.

In the 1990s, molecular phylogenetic analyses of DNA sequences produced a coherent scheme
showing arthropods as members of a superphylum labelled Ecdysozoa ("animals that moult"),
which contained nematodes, priapulids and tardigrades but excluded annelids. This was backed
up by studies of the anatomy and development of these animals, which showed that many of the
features that supported the Articulata hypothesis showed significant differences between annelids
and the earliest Panarthropods in their details, and some were hardly present at all in arthropods.
This hypothesis groups annelids with molluscs and brachiopods in another superphylum,
Lophotrochozoa.

If the Ecdysozoa hypothesis is correct, then segmentation of arthropods and annelids either has
evolved convergently or has been inherited from a much older ancestor and subsequently lost in
several other lineages, such as the non-arthropod members of the Ecdysozoa.[117][115]

Evolution of fossil arthropods

Further information: Deuteropoda

 Arthropod fossil phylogeny[118]
Arthropoda
giant lobopodians †

gilled lobopodians †

Radiodonta †
Deuteropoda
Chelicerata
Megacheira
†
Artiopoda †

Isoxyida †

Mandibulata

Summarized cladogram of the relationships

between extinct arthropod groups. For
more, see Deuteropoda.

Aside from the four major living groups (crustaceans, chelicerates, myriapods and hexapods), a
number of fossil forms, mostly from the early Cambrian period, are difficult to place
taxonomically, either from lack of obvious affinity to any of the main groups or from clear
affinity to several of them. Marrella was the first one to be recognized as significantly different
from the well-known groups.[44]

Modern interpretations of the basal, extinct stem-group of Arthropoda recognised the following
groups, from most basal to most crownward:[119][118]

 The "Giant" or "Siberiid Lobopodians", such as Jianshanopodia, Siberion and

Megadictyon, are the most basal grade in the total-group Arthropoda.
 The "Gilled Lobopodians", such as Kerygmachela, Pambdelurion and Opabinia, are the
second most basal grade.
 The Radiodonta, which traditionally known as anomalocaridids come in third position,
and are thought to be monophyletic.
 A possible "upper stem-group" assemblage of more uncertain position[118] but contained
within Deuteropoda:[119] the Fuxianhuiida, Megacheira, and multiple "bivalved forms"
including Isoxyida and Hymenocarina.

The Deuteropoda is a recently established clade uniting the crown-group (living) arthropods with
these possible "upper stem-group" fossils taxa.[119] The clade is defined by important changes to
the structure of the head region such as the appearance of a differentiated deutocerebral
appendage pair, which excludes more basal taxa like radiodonts and "gilled lobopodians".[119]
Controversies remain about the positions of various extinct arthropod groups. Some studies
recover Megacheira as closely related to chelicerates, while others recover them as outside the
group containing Chelicerate and Mandibulata as stem-group euarthropods.[120] The placement of
the Artiopoda (which contains the extinct trilobites and similar forms) is also a frequent subject
of dispute.[121] The main hypotheses position them in the clade Arachnomorpha with the
Chelicerates. However, one of the newer hypotheses is that the chelicerae have originated from
the same pair of appendages that evolved into antennae in the ancestors of Mandibulata, which
would place trilobites, which had antennae, closer to Mandibulata than Chelicerata, in the clade
Antennulata.[120][122] The fuxianhuiids, usually suggested to be stem-group arthropods, have been
suggested to be Mandibulates in some recent studies.[120] The Hymenocarina, a group of bivalved
arthropods, previously thought to have been stem-group members of the group, have been
demonstrated to be mandibulates based on the presence of mandibles.[118]

 Radiodonts, Opabiniids, Gilled Lobopodians and the more traditional Lobopodians are all
examples of basal stem-group arthropod lineages from the Cambrian

Anomalocaris
(Radiodonta)

Opabinia and Utaurora

(Opabiniidae)


 Mobulavermis
(Kerygmachelidae)

Facivermis
(Luolishaniidae)

 Marrellomorphs, megacherians, funxianhuiids and phosphatocopines are some examples

of Cambrian arthropods whose classification remains difficult

Marrella
(Marrellomorpha)

Leanchoilia
(Megacheira)


 Fuxianhuia
(Fuxianhuiida)

Dabashanella
(Phosphatocopina)

 Other examples of now extinct arthropod groups include

Acutiramus
(Eurypterida)

Apankura
(Euthycarcinoidea)


 Trimerus
(Artiopoda)

Concavicaris
(Thylacocephala)

Evolution and classification of living arthropods

See also: List of arthropod orders

The phylum Arthropoda is typically subdivided into four subphyla, of which one is extinct:[123]

1. Artiopods are an extinct group of formerly numerous marine animals that disappeared in
the Permian–Triassic extinction event, though they were in decline prior to this killing
blow, having been reduced to one order in the Late Devonian extinction. They contain
groups such as the trilobites.
2. Chelicerates comprise the marine sea spiders and horseshoe crabs, along with the
terrestrial arachnids such as mites, harvestmen, spiders, scorpions and related organisms
characterized by the presence of chelicerae, appendages just above/in front of the
mouthparts. Chelicerae appear in scorpions and horseshoe crabs as tiny claws that they
use in feeding, but those of spiders have developed as fangs that inject venom.
3. Myriapods comprise millipedes, centipedes, pauropods and symphylans, characterized
by having numerous body segments each of which bearing one or two pairs of legs (or in
a few cases being legless). All members are exclusively terrestrial.
4. Pancrustaceans comprise ostracods, barnacles, copepods, malacostracans,
cephalocaridans, branchiopods, remipedes and hexapods. Most groups are primarily
aquatic (two notable exceptions being woodlice and hexapods, which are both purely
terrestrial) and are characterized by having biramous appendages. The most abundant
group of pancrustaceans are the terrestrial hexapods, which comprise insects, diplurans,
springtails, and proturans, with six thoracic legs.

The phylogeny of the major extant arthropod groups has been an area of considerable interest
and dispute.[124] Recent studies strongly suggest that Crustacea, as traditionally defined, is
paraphyletic, with Hexapoda having evolved from within it,[125][126] so that Crustacea and
Hexapoda form a clade, Pancrustacea. The position of Myriapoda, Chelicerata and Pancrustacea
remains unclear as of April 2012. In some studies, Myriapoda is grouped with Chelicerata
(forming Myriochelata);[127][128] in other studies, Myriapoda is grouped with Pancrustacea
(forming Mandibulata),[125] or Myriapoda may be sister to Chelicerata plus Pancrustacea.[126]

The following cladogram shows the internal relationships between all the living classes of
arthropods as of the late 2010s,[129][130] as well as the estimated timing for some of the clades:[131]

Arthro Chelice Crusta

poda rata Pycnogonida ceans
Entogn
Euchelicerata aths
Xiphosura

Arachnida
Mandib Myriap
ulata oda Chilopoda
Progoneata Edafopoda
Symphyla

Pauropoda

Diplopoda
Pancrus Oligostr
tacea aca Ostracoda

Mystacocarida
Ichthyostraca
Branchiura

Pentastomida

Altocru Multicru Hexanauplia

stacea stacea Copepoda

Tantulocarida

Thecostraca

Malacostraca
Allotrioc
arida Cephalocarida
Athalass
ocarida Branchiopoda
493 mya Labioc
arida
 Subphyla Classes Members Example species

sea spiders
Pycnogonida
horseshoe crabs
Chelicerata Xiphosura
harvestmen, solifuges, mites, scorpions,
Arachnida
spiders, ticks etc.
Platycryptus undatus
(Arachnida, Araneae)

pseudocentipedes
Symphyla hexameroceratans, tetrameroceratans
Pauropoda bristle millipedes, pill millipedes, flat-
Myriapoda
Diplopoda backed millipedes, etc.
Chilopoda scutigeromorphs, lithobiomorphs,
Scolopendromorphs, etc. Archispirostreptus gigas
(Diplopoda,
Spirostreptida)
Crustacea Ostracoda seed shrimp
Mystacocarida Mystacocaridans
Pentastomida tongue worms
Branchiura fish lice
Thecostraca barnacles, etc.
Copepoda calanoids, cyclopoids, misophrioids,
Malacostraca siphonostomatoids, etc. Ocypode ceratophthalma
Cephalocarida mantis shrimp, skeleton shrimp, (Malacostraca, Decapoda)
Branchiopoda woodlice, shrimp, crabs, lobsters, krill,
Remipedia etc.
horseshoe shrimp
fairy shrimp, tadpole shrimp, water fleas,
clam shrimp
 remipedes

Insecta insects
Hexapoda
Entognatha springtails, etc.

Saturnia pavonia
(Insecta, Lepidoptera)

Interaction with humans

Main article: Arthropods in culture
See also: Insects as food

Insects and scorpions on sale in a food stall in Bangkok,

Thailand

Crustaceans such as crabs, lobsters, crayfish, shrimp, and prawns have long been part of human
cuisine, and are now raised commercially.[132] Insects and their grubs are at least as nutritious as
meat, and are eaten both raw and cooked in many cultures, though not most European, Hindu,
and Islamic cultures.[133][134] Cooked tarantulas are considered a delicacy in Cambodia,[135][136][137]
and by the Piaroa Indians of southern Venezuela, after the highly irritant hairs – the spider's main
defense system – are removed.[138] Humans also unintentionally eat arthropods in other foods,[139]
and food safety regulations lay down acceptable contamination levels for different kinds of food
material.[Note 4][Note 5] The intentional cultivation of arthropods and other small animals for human
food, referred to as minilivestock, is now emerging in animal husbandry as an ecologically sound
concept.[143] Commercial butterfly breeding provides Lepidoptera stock to butterfly
conservatories, educational exhibits, schools, research facilities, and cultural events.

However, the greatest contribution of arthropods to human food supply is by pollination: a 2008
study examined the 100 crops that FAO lists as grown for food, and estimated pollination's
economic value as €153 billion, or 9.5 per cent of the value of world agricultural production used
for human food in 2005.[144] Besides pollinating, bees produce honey, which is the basis of a
rapidly growing industry and international trade.[145]

The red dye cochineal, produced from a Central American species of insect, was economically
important to the Aztecs and Mayans.[146] While the region was under Spanish control, it became
Mexico's second most-lucrative export,[147] and is now regaining some of the ground it lost to
synthetic competitors.[148] Shellac, a resin secreted by a species of insect native to southern Asia,
was historically used in great quantities for many applications in which it has mostly been
replaced by synthetic resins, but it is still used in woodworking and as a food additive. The blood
of horseshoe crabs contains a clotting agent, Limulus Amebocyte Lysate, which is now used to
test that antibiotics and kidney machines are free of dangerous bacteria, and to detect spinal
meningitis and some cancers.[149] Forensic entomology uses evidence provided by arthropods to
establish the time and sometimes the place of death of a human, and in some cases the cause. [150]
Recently insects have also gained attention as potential sources of drugs and other medicinal
substances.[151]

The relative simplicity of the arthropods' body plan, allowing them to move on a variety of
surfaces both on land and in water, have made them useful as models for robotics. The
redundancy provided by segments allows arthropods and biomimetic robots to move normally
even with damaged or lost appendages.[152][153]

Diseases transmitted by insects

[154]
Disease Insect Cases per year Deaths per year
Malaria Anopheles mosquito 267 M 1 to 2 M
Dengue fever Aedes mosquito ? ?
Yellow fever Aedes mosquito 4,432 1,177
Filariasis Culex mosquito 250 M unknown

Although arthropods are the most numerous phylum on Earth, and thousands of arthropod
species are venomous, they inflict relatively few serious bites and stings on humans. Far more
serious are the effects on humans of diseases like malaria carried by blood-sucking insects. Other
blood-sucking insects infect livestock with diseases that kill many animals and greatly reduce the
usefulness of others.[154] Ticks can cause tick paralysis and several parasite-borne diseases in
humans.[155] A few of the closely related mites also infest humans, causing intense itching,[156] and
others cause allergic diseases, including hay fever, asthma, and eczema.[157]

Many species of arthropods, principally insects but also mites, are agricultural and forest pests.
[158][159]
The mite Varroa destructor has become the largest single problem faced by beekeepers
worldwide.[160] Efforts to control arthropod pests by large-scale use of pesticides have caused
long-term effects on human health and on biodiversity.[161] Increasing arthropod resistance to
pesticides has led to the development of integrated pest management using a wide range of
measures including biological control.[158] Predatory mites may be useful in controlling some
mite pests.[162][163]

See also
 Dorsal lobe
 Invertebrate paleontology
 Minibeasts

Notes
 1.

 The Museum of New Zealand notes that "in everyday conversation", bug "refers to land
arthropods with at least six legs, such as insects, spiders, and centipedes".[21] In a chapter on
"Bugs That Are Not Insects", entomologist Gilbert Walbauer specifies centipedes, millipedes,
arachnids (spiders, daddy longlegs, scorpions, mites, chiggers and ticks) as well as the few
terrestrial crustaceans (sowbugs and pillbugs),[22] but argues that "including legless creatures such
as worms, slugs, and snails among the bugs stretches the word too much".[23]
  "It would be too bad if the question of head segmentation ever should be finally settled; it
has been for so long such fertile ground for theorizing that arthropodists would miss it as a field
for mental exercise."[46]
  The fossil was originally named Eotarbus but was renamed when it was realized that a
Carboniferous arachnid had already been named Eotarbus.[96]
  For a mention of insect contamination in an international food quality standard, see
sections 3.1.2 and 3.1.3 of Codex 152 of 1985 of the Codex Alimentarius[140]

5.  For examples of quantified acceptable insect contamination levels in food see the last
entry (on "Wheat Flour") and the definition of "Extraneous material" in Codex
Alimentarius,[141] and the standards published by the FDA.[142]

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External links
Arthropod at Wikipedia's sister projects

 Definitions from Wiktionary

 Media from Commons
 Quotations from Wikiquote
 Texts from Wikisource
 Textbooks from Wikibooks
 Resources from Wikiversity
 Taxa from Wikispecies
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 "Arthropod" at the Encyclopedia of Life

 Venomous Arthropods chapter in United States Environmental Protection Agency and
University of Florida/Institute of Food and Agricultural Sciences National Public Health
Pesticide Applicator Training Manual
 Arthropods – Arthropoda Insect Life Forms


 v
 t
 e

Extant animal phyla

 v
 t
 e
Extant Arthropoda classes by subphylum

 v
 t
 e

Extant life phyla/divisions by domain

Taxon identifiers
 Wikidata: Q1360
 Wikispecies: Arthropoda
 ADW: Arthropoda
 AFD: Arthropoda
 BOLD: 20
 BugGuide: 3
 EoL: 164
 EPPO: 1ARTHP
 Fauna Europaea: 2
 Fauna Europaea (new): a068518a-257a-4dbb-8425-b6faaef7d56b
 Fossilworks: 18891
Arthropoda
 GBIF: 54
 iNaturalist: 47120
 IRMNG: 235
 ITIS: 82696
 MilliBase: 1065
 NCBI: 6656
 NZOR: 3ccd1beb-10c5-4af0-861d-4030f74d80a8
 Open Tree of Life: 632179
 PPE: arthropoda
 WoRMS: 1065

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Categories:

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 Extant Cambrian first appearances

 This page was last edited on 15 March 2024, at 09:22 (UTC).

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