Arthropod
Encyclopedia
An arthropod is an invertebrate
animal
having an exoskeleton
(external skeleton
), a segmented body, and jointed appendage
s. Arthropods are members of the phylum
Arthropoda (from Greek
, "joint
", and "foot
", which together mean "jointed feet"), and include the insect
s, arachnid
s, crustacean
s, and others. Arthropods are characterized by their jointed limbs and cuticle
s, which are mainly made of α-chitin
; the cuticles of crustaceans are also biomineralized with calcium carbonate
. The rigid cuticle inhibits growth, so arthropods replace it periodically by molting. The arthropod body plan
consists of repeated segments, each with a pair of appendage
s. It is so versatile that they have been compared to Swiss Army knives
, and it has enabled them to become the most species-rich members of all ecological guilds
in most environments. They have over a million described species, making up more than 80% of all described living animal species, and are one of only two animal groups that are very successful in dry environments – the other being the amniote
s. They range in size from microscopic plankton
up to forms a few meters long.
Arthropods' primary internal cavity is a hemocoel, which accommodates their internal organs
and through which their blood
circulates; they have open circulatory systems. 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 brain
s 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.
Their vision relies on various combinations of compound eyes and pigment-pit ocelli: 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 spider
s 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 seta
e (bristles) that project through their cuticles.
Arthropods' methods of reproduction and development are diverse; all terrestrial species use internal fertilization
, but this is often 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, but scorpion
s give birth to live young after the eggs have hatched inside the mother. Arthropod hatchlings vary from miniature adults to grubs and caterpillar
s 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 scorpions.
The versatility of the arthropod modular body plan has made it difficult for zoologists and paleontologists to classify them and work out their evolutionary ancestry, which dates back to the Cambrian
period. From the late 1950s to late 1970s, it was thought that arthropods were polyphyletic, that is, there was no single arthropod ancestor. Now they are generally regarded as monophyletic. Historically, the closest evolutionary relatives of arthropods were considered to be annelid
worms, as both groups have segmented bodies. This hypothesis is by now largely rejected, with annelids and molluscs
forming the superphylum Lophotrochozoa
. Many analyses support a placement of arthropods with cycloneuralia
ns (or their constituent clades) in a superphylum Ecdysozoa
. Overall however, the basal relationships of Metazoa are not yet well resolved. Likewise, the relationships between various arthropod groups are still actively debated.
Arthropods contribute to the human food supply both directly as food, and more importantly as pollinators of crops. Some specific species are known to spread severe disease to humans, livestock
, and crops.
s with segmented
bodies and jointed limbs. The limbs form part of an exoskeleton
, which is mainly made of α-chitin
, a derivative of glucose
. One other group of animals, the tetrapod
s, has jointed limbs, but tetrapods are vertebrate
s and therefore have endoskeleton
s.
, and account for over 80% of all known living animal species. Another study estimates that there are between 5 to 10 million extant arthropod species, both described and yet to be described. Estimating the total number of living species is extremely difficult because it often depends on a series of assumptions in order to scale up from counts at specific locations to estimates for 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.
They are important members of marine, freshwater, land and air ecosystem
s, and are one of only two major animal groups that have adapted to life in dry environments; the other is amniote
s, whose living members are reptile
s, bird
s and mammal
s. One arthropod sub-group, insect
s, is the most species-rich member of all ecological guilds
(ways of making a living) in land and fresh-water environments. The lightest insects weigh less than 25 micrograms (millionths of a gram), while the heaviest weigh over 70 grams (2.5 oz). Some living crustacean
s are much larger; for example, the legs of the Japanese spider crab
may span up to 4 metres (13.1 ft).
s 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; The three-part appearance of many insect
bodies and the two-part appearance of spider
s is a result of this grouping; in fact there are no external signs of segmentation in mite
s. Arthropods also have two body elements that are not part of this serially repeated pattern of segments, an acron
at the front, ahead of the mouth, and a telson
at the rear, behind the anus
. The eyes are mounted on the acron.
The original structure of arthropod appendages was probably biramous, with the upper branch acting as a gill
while the lower branch was used for walking. In some segments of all known arthropods the appendages have been modified, for example to form gills, mouth-parts, antennae
for collecting information, or claws for grasping; arthropods are "like Swiss Army knives
, each equipped with a unique set of specialized tools." In many arthropods, appendages have vanished from some regions of the body, and it is particularly common for abdominal appendages to have disappeared or be highly modified.
The most conspicuous specialization of segments is in the head. The four major groups of arthropods – Chelicerata
(includes spider
s and scorpion
s), Crustacea
(shrimp
s, lobster
s, crab
s, etc.), Tracheata (arthropods that breathe via channels into their bodies; includes insect
s and myriapods), and the extinct trilobite
s – have heads formed of various combinations of segments, with appendages that are missing or specialized in different ways. 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. 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
". In 1960 R. E. Snodgrass even hoped it would not be solved, as trying to work out solutions was so much fun.
, a non-cellular material secreted by the epidermis
. Their cuticles vary in the details of their structure, but generally consist of three main layers: the epicuticle, a thin outer wax
y coat that moisture-proofs the other layers and gives them some protection; the exocuticle, which consists of chitin
and chemically hardened protein
s; and the endocuticle, which consists of chitin and unhardened proteins. The exocuticle and endocuticle together are known as the procuticle. 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.
The exoskeletons of most aquatic crustacean
s 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. Biomineralization generally affects the exocuticle and the outer part of the endocuticle. Two recent hypotheses about the evolution of biomineralization in arthropods and other groups of animals propose that it provides tougher defensive armor, and that it allows animals to grow larger and stronger by providing more rigid skeletons; and in either case a mineral-organic composite
exoskeleton is cheaper to build than an all-organic one of comparable strength.
The cuticle can have seta
e (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.
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; for example, all spiders extend their legs hydraulically and can generate pressures up to eight times their resting level.
In the initial phase of molting, the animal stops feeding and its epidermis releases molting fluid, a mixture of enzyme
s 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.
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. Molting may be responsible for 80 to 90% of all arthropod deaths.
, 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.
Arthropods have open circulatory system
s, although 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, but few chelicerates and tracheates, use respiratory pigment
s to assist oxygen transport. The most common respiratory pigment in arthropods is copper
-based hemocyanin
; this is used by many crustaceans and a few centipede
s. A few crustaceans and insects use iron-based hemoglobin
, the respiratory pigment used by vertebrate
s. As with other invertebrates and unlike among vertebrates, the respiratory pigments of those arthropods that have them are generally dissolved in the blood and rarely enclosed in corpuscles
.
The heart is typically 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 ligament
s or by small muscle
s, 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.
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 lung
s. 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 arachnid
s.
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 commissure
s (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").
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. 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. 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
. Most aquatic arthropods and some terrestrial ones also have organs called nephridia ("little kidney
s"), which extract other wastes for excretion as urine
.
s 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 seta
e, 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 eardrum
s, 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.
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 spider
s are pigment-cup ocelli that are capable of forming images, and those of jumping spider
s can rotate to track prey.
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
. Compound eyes have a wide field of view, and can detect fast movement and, in some cases, the polarization of light. 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 centimetres (7.9 in) are most important to most arthropods. 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.
Most arthropods lack balance and acceleration
sensors, and rely on their eyes to tell them which way is up. The self-righting behavior of cockroach
es is triggered when pressure sensors on the underside of the feet report no pressure. However many malacostracan crustacean
s have statocyst
s, which provide the same sort of information as the balance and motion sensors of the vertebrate inner ear
.
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.
A few arthropods, such as barnacle
s, are hermaphroditic, that is, each can have the organs of both sex
es. However, individuals of most species remain of one sex all their lives. A few species of insect
s and crustacean
s can reproduce by parthenogenesis
, for example, without mating, 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. Aquatic
arthropods may breed by external fertilization
, as for example frog
s also do, 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, as unprotected sperm
and ova would not survive long in these environments. In a few cases the sperm transfer is direct from the male's penis
to the female's oviduct
, but it is more often indirect. Some crustaceans and spiders use modified appendages to transfer the sperm to the female. On the other hand, many male terrestrial
arthropods produce spermatophore
s, 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.
Most arthropods lay eggs, but scorpions are viviparous: they produce live young after the eggs have hatched inside the mother, and are noted for prolonged maternal care. 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 molt. Many insects hatch as grubs or caterpillar
s, 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. Dragonfly
larvae have the typical cuticles and jointed limbs of arthropods but are flightless water-breathers with extendable jaws. Crustaceans commonly hatch as tiny nauplius larvae that have only three segments and pairs of appendages.
(armor plate) and bearing a pair of biramous limbs. Whether the ancestral limb was uniramous or biramous is far from a settled debate, though.
This Ur-arthropod had a ventral mouth, pre-oral antennae and dorsal eyes at the front of the body. It was a non-discriminatory sediment
feeder, processing whatever sediment came its way for food.
It has been proposed that the Ediacaran
animals Parvancorina
and Spriggina
, from around 555 Mya, were arthropods. Small arthropods with bivalve-like shells have been found in Early Cambrian fossil beds dating in China. The earliest Cambrian trilobite
fossils are about 530 million years old, but the class was already quite diverse and worldwide, suggesting that they had been around for quite some time. Re-examination in the 1970s of the Burgess Shale
fossils from about 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
. A fossil of Marrella from the Burgess Shale has provided the earliest clear evidence of molting.
The earliest fossil crustacean
s date from about in the Cambrian
, and fossil shrimp
from about apparently formed a tight-knit procession across the seabed. Crustacean fossils are common from the Ordovician
period onwards. They have remained almost entirely aquatic, possibly because they never developed excretory system
s that conserve water.
Arthropods provide the earliest identifiable fossils of land animals, from about in the Late Silurian
, and terrestrial tracks from about appear to have been made by arthropods. Arthropods were well pre-adapted
to colonize land, because their existing jointed exoskeletons provided protection against desiccation, support against gravity and a means of locomotion that was not dependent on water. Around the same time the aquatic, scorpion-like eurypterid
s became the largest ever arthropods, some as long as 2.5 metres (8.2 ft).
The oldest known arachnid
is the trigonotarbid Palaeotarbus jerami, from about in the Silurian
period. Attercopus
fimbriunguis, from in the Devonian
period, bears the earliest known silk-producing spigots, but its lack of spinnerets means it was not one of the true spider
s, which first appear in the Late Carboniferous
over . The Jurassic
and Cretaceous
periods provide a large number of fossil spiders, including representatives of many modern families. Fossils of aquatic scorpion
s with gill
s appear in the Silurian and Devonian
periods, and the earliest fossil of an air-breathing scorpion with book lung
s dates from the Early Carboniferous period.
The oldest definitive insect fossil is the Devonian
Rhyniognatha hirsti, dated at , but its mandibles are of a type found only in winged insects, which suggests that the earliest insects appeared in the Silurian period. The Mazon Creek lagerstätten
from the Late Carboniferous, about , include about 200 species, some gigantic by modern standards, and indicate that insects had occupied their main modern ecological niche
s as herbivore
s, detritivore
s and insectivore
s. 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 Mid Cenozoic
.
From 1952 to 1977, Sidnie Manton
and others argued that arthropods are polyphyletic, in other words, 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 spider
s and scorpion
s; the crustacean
s; and the uniramia
, consisting of onychophorans, myriapods and hexapods
. These arguments usually bypassed trilobite
s, 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 exoskeleton
s; 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.
including Hallucigenia
and Microdictyon
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Simplified summary of Budd's "broad-scale" cladogram (1996)Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...
animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...
having an exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
(external skeleton
Skeleton
The skeleton is the body part that forms the supporting structure of an organism. There are two different skeletal types: the exoskeleton, which is the stable outer shell of an organism, and the endoskeleton, which forms the support structure inside the body.In a figurative sense, skeleton can...
), a segmented body, and jointed appendage
Appendage
In invertebrate biology, an appendage is an external body part, or natural prolongation, that protrudes from an organism's body . It is a general term that covers any of the homologous body parts that may extend from a body segment...
s. Arthropods are members of the phylum
Phylum
In biology, a phylum The term was coined by Georges Cuvier from Greek φῦλον phylon, "race, stock," related to φυλή phyle, "tribe, clan." is a taxonomic rank below kingdom and above class. "Phylum" is equivalent to the botanical term division....
Arthropoda (from Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
, "joint
Joint
A joint is the location at which two or more bones make contact. They are constructed to allow movement and provide mechanical support, and are classified structurally and functionally.-Classification:...
", and "foot
Foot
The foot is an anatomical structure found in many vertebrates. It is the terminal portion of a limb which bears weight and allows locomotion. In many animals with feet, the foot is a separate organ at the terminal part of the leg made up of one or more segments or bones, generally including claws...
", which together mean "jointed feet"), and include the insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s, arachnid
Arachnid
Arachnids are a class of joint-legged invertebrate animals in the subphylum Chelicerata. All arachnids have eight legs, although in some species the front pair may convert to a sensory function. The term is derived from the Greek words , meaning "spider".Almost all extant arachnids are terrestrial...
s, crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s, and others. Arthropods are characterized by their jointed limbs and cuticle
Cuticle
A cuticle , or cuticula, is a term used for any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticles" are non-homologous; differing in their origin, structure, function, and chemical composition...
s, which are mainly made of α-chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
; the cuticles of crustaceans are also biomineralized with calcium carbonate
Calcium carbonate
Calcium carbonate is a chemical compound with the formula CaCO3. It is a common substance found in rocks in all parts of the world, and is the main component of shells of marine organisms, snails, coal balls, pearls, and eggshells. Calcium carbonate is the active ingredient in agricultural lime,...
. The rigid cuticle inhibits growth, so arthropods replace it periodically by molting. The arthropod body plan
Body plan
A body plan is the blueprint for the way the body of an organism is laid out. An organism's symmetry, its number of body segments and number of limbs are all aspects of its body plan...
consists of repeated segments, each with a pair of appendage
Appendage
In invertebrate biology, an appendage is an external body part, or natural prolongation, that protrudes from an organism's body . It is a general term that covers any of the homologous body parts that may extend from a body segment...
s. It is so versatile that they have been compared to Swiss Army knives
Swiss Army knife
The Swiss Army knife is a brand of pocket knife or multi-tool manufactured by Victorinox AG and Wenger SA. The term "Swiss Army knife" was coined by US soldiers after World War II due to the difficulty they had in pronouncing the German name....
, and it has enabled them to become the most species-rich members of all ecological guilds
Guild (ecology)
A guild is any group of species that exploit the same resources, often in related ways. As can be seen from the list of examples below, it does not follow that the species within a guild occupy the same, or even similar, ecological niches...
in most environments. They have over a million described species, making up more than 80% of all described living animal species, and are one of only two animal groups that are very successful in dry environments – the other being the amniote
Amniote
The amniotes are a group of tetrapods that have a terrestrially adapted egg. They include synapsids and sauropsids , as well as their fossil ancestors. Amniote embryos, whether laid as eggs or carried by the female, are protected and aided by several extensive membranes...
s. They range in size from microscopic plankton
Plankton
Plankton are any drifting organisms that inhabit the pelagic zone of oceans, seas, or bodies of fresh water. That is, plankton are defined by their ecological niche rather than phylogenetic or taxonomic classification...
up to forms a few meters long.
Arthropods' primary internal cavity is a hemocoel, which accommodates their internal organs
Organ (anatomy)
In biology, an organ is a collection of tissues joined in structural unit to serve a common function. Usually there is a main tissue and sporadic tissues . The main tissue is the one that is unique for the specific organ. For example, main tissue in the heart is the myocardium, while sporadic are...
and through which their blood
Blood
Blood is a specialized bodily fluid in animals that delivers necessary substances such as nutrients and oxygen to the cells and transports metabolic waste products away from those same cells....
circulates; they have open circulatory systems. Like their exteriors, the internal organs of arthropods are generally built of repeated segments. Their nervous system
Nervous system
The nervous system is an organ system containing a network of specialized cells called neurons that coordinate the actions of an animal and transmit signals between different parts of its body. In most animals the nervous system consists of two parts, central and peripheral. The central nervous...
is "ladder-like", with paired ventral nerve cords
Ventral nerve cord
The ventral nerve cord makes up the nervous system of some phyla of the invertebrates, particularly within the nematodes, annelids and the arthropods. It usually consists of cerebral ganglia anteriorly with the nerve cords running down the ventral plane of the organism...
running through all segments and forming paired ganglia in each segment. Their heads are formed by fusion of varying numbers of segments, and their brain
Brain
The brain is the center of the nervous system in all vertebrate and most invertebrate animals—only a few primitive invertebrates such as sponges, jellyfish, sea squirts and starfishes do not have one. It is located in the head, usually close to primary sensory apparatus such as vision, hearing,...
s are formed by fusion of the ganglia of these segments and encircle the esophagus
Esophagus
The esophagus is an organ in vertebrates which consists of a muscular tube through which food passes from the pharynx to the stomach. During swallowing, food passes from the mouth through the pharynx into the esophagus and travels via peristalsis to the stomach...
. The respiratory
Respiratory system
The respiratory system is the anatomical system of an organism that introduces respiratory gases to the interior and performs gas exchange. In humans and other mammals, the anatomical features of the respiratory system include airways, lungs, and the respiratory muscles...
and excretory
Excretion
Excretion is the process by which waste products of metabolism and other non-useful materials are eliminated from an organism. This is primarily carried out by the lungs, kidneys and skin. This is in contrast with secretion, where the substance may have specific tasks after leaving the cell...
systems of arthropods vary, depending as much on their environment as on the subphylum
Subphylum
In life, a subphylum is a taxonomic rank intermediate between phylum and superclass. The rank of subdivision in plants and fungi is equivalent to subphylum.Not all phyla are divided into subphyla...
to which they belong.
Their vision relies on various combinations of compound eyes and pigment-pit ocelli: 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 spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s 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 seta
Seta
Seta is a biological term derived from the Latin word for "bristle". It refers to a number of different bristle- or hair-like structures on living organisms.-Animal setae:In zoology, most "setae" occur in invertebrates....
e (bristles) that project through their cuticles.
Arthropods' methods of reproduction and development are diverse; all terrestrial species use internal fertilization
Internal fertilization
In mammals, internal fertilization is done through copulation, which involves the insertion of the penis into the vagina. Some other higher vertebrate animals reproduce internally, but their fertilization is cloacal.The union of spermatozoa of the parent organism. At some point, the growing egg or...
, but this is often 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
External fertilization
External fertilization is a form of fertilization in which a sperm cell is united with an egg cell external to the bodies of the reproducing individuals. In contrast, internal fertilization takes place inside the female after insemination through copulation....
. Almost all arthropods lay eggs, but scorpion
Scorpion
Scorpions are predatory arthropod animals of the order Scorpiones within the class Arachnida. They have eight legs and are easily recognized by the pair of grasping claws and the narrow, segmented tail, often carried in a characteristic forward curve over the back, ending with a venomous stinger...
s give birth to live young after the eggs have hatched inside the mother. Arthropod hatchlings vary from miniature adults to grubs and caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...
s that lack jointed limbs and eventually undergo a total metamorphosis
Metamorphosis
Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation...
to produce the adult form. The level of maternal care for hatchlings varies from nonexistent to the prolonged care provided by scorpions.
The versatility of the arthropod modular body plan has made it difficult for zoologists and paleontologists to classify them and work out their evolutionary ancestry, which dates back to the Cambrian
Cambrian
The Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...
period. From the late 1950s to late 1970s, it was thought that arthropods were polyphyletic, that is, there was no single arthropod ancestor. Now they are generally regarded as monophyletic. Historically, the closest evolutionary relatives of arthropods were considered to be annelid
Annelid
The annelids , formally called Annelida , are a large phylum of segmented worms, with over 17,000 modern species including ragworms, earthworms and leeches...
worms, as both groups have segmented bodies. This hypothesis is by now largely rejected, with annelids and molluscs
Mollusca
The Mollusca , common name molluscs or mollusksSpelled mollusks in the USA, see reasons given in Rosenberg's ; for the spelling mollusc see the reasons given by , is a large phylum of invertebrate animals. There are around 85,000 recognized extant species of molluscs. Mollusca is the largest...
forming the superphylum Lophotrochozoa
Lophotrochozoa
The Lophotrochozoa are a major grouping of protostome animals. The taxon was discovered based on molecular data. Molecular evidence such as a result of studies of the evolution of small-subunit ribosomal RNA supports the monophyly of the phyla listed in the infobox shown at right.-Terminology:The...
. Many analyses support a placement of arthropods with cycloneuralia
Cycloneuralia
Cycloneuralia is a clade of ecdysozoan animals including the Scalidophora and the Nematoida . Its sister group is the panarthropoda....
ns (or their constituent clades) in a superphylum Ecdysozoa
Ecdysozoa
Ecdysozoa is a group of protostome animals, including Arthropoda , Nematoda, and several smaller phyla. They were first defined by Aguinaldo et al. in 1997, based mainly on trees constructed using 18S ribosomal RNA genes...
. Overall however, the basal relationships of Metazoa are not yet well resolved. Likewise, the relationships between various arthropod groups are still actively debated.
Arthropods contribute to the human food supply both directly as food, and more importantly as pollinators of crops. Some specific species are known to spread severe disease to humans, livestock
Livestock
Livestock refers to one or more domesticated animals raised in an agricultural setting to produce commodities such as food, fiber and labor. The term "livestock" as used in this article does not include poultry or farmed fish; however the inclusion of these, especially poultry, within the meaning...
, and crops.
Description
Arthropods are invertebrateInvertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...
s with segmented
Segmentation (biology)
Segmentation in biology refers to either a type of gastrointestinal motility or the division of some animal and plant body plans into a series of repetitive segments. This article will focus on the segmentation of animal body plans, specifically using the examples of the phyla Arthropoda,...
bodies and jointed limbs. The limbs form part of an exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
, which is mainly made of α-chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
, a derivative of glucose
Glucose
Glucose is a simple sugar and an important carbohydrate in biology. Cells use it as the primary source of energy and a metabolic intermediate...
. One other group of animals, the tetrapod
Tetrapod
Tetrapods are vertebrate animals having four limbs. Amphibians, reptiles, birds and mammals are all tetrapods; even snakes and other limbless reptiles and amphibians are tetrapods by descent. The earliest tetrapods evolved from the lobe-finned fishes in the Devonian...
s, has jointed limbs, but tetrapods are vertebrate
Vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...
s and therefore have endoskeleton
Endoskeleton
An endoskeleton is an internal support structure of an animal, composed of mineralized tissue. Endoskeleton develops within the skin or in the deeper body tissues. The vertebrate is basically an endoskeleton made up of two types of tissues . During early embryonic development the endoskeleton is...
s.
Diversity
One estimate indicates that arthropods have 1,170,000 described speciesSpecies description
A species description or type description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species which have been described previously, or are...
, and account for over 80% of all known living animal species. Another study estimates that there are between 5 to 10 million extant arthropod species, both described and yet to be described. Estimating the total number of living species is extremely difficult because it often depends on a series of assumptions in order to scale up from counts at specific locations to estimates for 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.
They are important members of marine, freshwater, land and air ecosystem
Ecosystem
An ecosystem is a biological environment consisting of all the organisms living in a particular area, as well as all the nonliving , physical components of the environment with which the organisms interact, such as air, soil, water and sunlight....
s, and are one of only two major animal groups that have adapted to life in dry environments; the other is amniote
Amniote
The amniotes are a group of tetrapods that have a terrestrially adapted egg. They include synapsids and sauropsids , as well as their fossil ancestors. Amniote embryos, whether laid as eggs or carried by the female, are protected and aided by several extensive membranes...
s, whose living members are reptile
Reptile
Reptiles are members of a class of air-breathing, ectothermic vertebrates which are characterized by laying shelled eggs , and having skin covered in scales and/or scutes. They are tetrapods, either having four limbs or being descended from four-limbed ancestors...
s, bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...
s and mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...
s. One arthropod sub-group, insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s, is the most species-rich member of all ecological guilds
Guild (ecology)
A guild is any group of species that exploit the same resources, often in related ways. As can be seen from the list of examples below, it does not follow that the species within a guild occupy the same, or even similar, ecological niches...
(ways of making a living) in land and fresh-water environments. The lightest insects weigh less than 25 micrograms (millionths of a gram), while the heaviest weigh over 70 grams (2.5 oz). Some living crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s are much larger; for example, the legs of the Japanese spider crab
Japanese spider crab
The , Macrocheira kaempferi, is a species of marine crab that lives in the waters around Japan. It has the largest leg span of any arthropod, reaching up to and weighing up to . It is the subject of small-scale fishery.-Description:...
may span up to 4 metres (13.1 ft).
Segmentation
The embryoEmbryo
An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...
s of all arthropods are segmented, built from a series of repeated modules. The last common ancestor
Most recent common ancestor
In genetics, the most recent common ancestor of any set of organisms is the most recent individual from which all organisms in the group are directly descended...
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
Tagma (biology)
In invertebrate biology, a tagma is a specialized grouping of arthropod segments, such as the head, the thorax, and the abdomen with a common function. The segments of a tagma may be either fused or moveable.-Tagmata:...
in which segments and their limbs are specialized in various ways; The three-part appearance of many insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
bodies and the two-part appearance of spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s is a result of this grouping; in fact there are no external signs of segmentation in mite
Mite
Mites, along with ticks, are small arthropods belonging to the subclass Acari and the class Arachnida. The scientific discipline devoted to the study of ticks and mites is called acarology.-Diversity and systematics:...
s. Arthropods also have two body elements that are not part of this serially repeated pattern of segments, an acron
Prostomium
Prostomium is the first body segment in annelid worms. It is in front of the mouth, being usually a small shelf- or lip-like extension over the dorsal side of the mouth. It sometimes bears antennae and eyes. It often functions like a kind of overlip when the animal is feeding...
at the front, ahead of the mouth, and a telson
Telson
The telson is the last division of the body of a crustacean. It is not considered a true segment because it does not arise in the embryo from teloblast areas as do real segments. It never carries any appendages, but a forked "tail" called the caudal furca is often present. Together with the...
at the rear, behind the anus
Anus
The anus is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...
. The eyes are mounted on the acron.
The original structure of arthropod appendages was probably biramous, with the upper branch acting as a gill
Gill
A gill is a respiratory organ found in many aquatic organisms that extracts dissolved oxygen from water, afterward excreting carbon dioxide. The gills of some species such as hermit crabs have adapted to allow respiration on land provided they are kept moist...
while the lower branch was used for walking. In some segments of all known arthropods the appendages have been modified, for example to form gills, mouth-parts, antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
for collecting information, or claws for grasping; arthropods are "like Swiss Army knives
Swiss Army knife
The Swiss Army knife is a brand of pocket knife or multi-tool manufactured by Victorinox AG and Wenger SA. The term "Swiss Army knife" was coined by US soldiers after World War II due to the difficulty they had in pronouncing the German name....
, each equipped with a unique set of specialized tools." In many arthropods, appendages have vanished from some regions of the body, and it is particularly common for abdominal appendages to have disappeared or be highly modified.
The most conspicuous specialization of segments is in the head. The four major groups of arthropods – Chelicerata
Chelicerata
The subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda, and includes horseshoe crabs, scorpions, spiders and mites...
(includes spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s and scorpion
Scorpion
Scorpions are predatory arthropod animals of the order Scorpiones within the class Arachnida. They have eight legs and are easily recognized by the pair of grasping claws and the narrow, segmented tail, often carried in a characteristic forward curve over the back, ending with a venomous stinger...
s), Crustacea
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
(shrimp
Shrimp
Shrimp are swimming, decapod crustaceans classified in the infraorder Caridea, found widely around the world in both fresh and salt water. Adult shrimp are filter feeding benthic animals living close to the bottom. They can live in schools and can swim rapidly backwards. Shrimp are an important...
s, lobster
Lobster
Clawed lobsters comprise a family of large marine crustaceans. Highly prized as seafood, lobsters are economically important, and are often one of the most profitable commodities in coastal areas they populate.Though several groups of crustaceans are known as lobsters, the clawed lobsters are most...
s, crab
Crab
True crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...
s, etc.), Tracheata (arthropods that breathe via channels into their bodies; includes insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s and myriapods), and the extinct trilobite
Trilobite
Trilobites are a well-known fossil group of extinct marine arthropods that form the class Trilobita. The first appearance of trilobites in the fossil record defines the base of the Atdabanian stage of the Early Cambrian period , and they flourished throughout the lower Paleozoic era before...
s – have heads formed of various combinations of segments, with appendages that are missing or specialized in different ways. 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. 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
Arthropod head problem
The arthropod head problem is a long-standing zoological dispute concerning the segmental composition of the heads of the various arthropod groups, and how they are evolutionarily related to each other...
". In 1960 R. E. Snodgrass even hoped it would not be solved, as trying to work out solutions was so much fun.
Exoskeleton
Arthropod exoskeletons are made of cuticleCuticle
A cuticle , or cuticula, is a term used for any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticles" are non-homologous; differing in their origin, structure, function, and chemical composition...
, a non-cellular material secreted by the epidermis
Epidermis (zoology)
The Epidermis is an epithelium that covers the body of an eumetazoan . Eumetazoa have a cavity lined with a similar epithelium, the gastrodermis, which forms a boundary with the epidermis at the mouth.Sponges have no epithelium, and therefore no epidermis or gastrodermis...
. Their cuticles vary in the details of their structure, but generally consist of three main layers: the epicuticle, a thin outer wax
Wax
thumb|right|[[Cetyl palmitate]], a typical wax ester.Wax refers to a class of chemical compounds that are plastic near ambient temperatures. Characteristically, they melt above 45 °C to give a low viscosity liquid. Waxes are insoluble in water but soluble in organic, nonpolar solvents...
y coat that moisture-proofs the other layers and gives them some protection; the exocuticle, which consists of chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
and chemically hardened protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
s; and the endocuticle, which consists of chitin and unhardened proteins. The exocuticle and endocuticle together are known as the procuticle. 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.
The exoskeletons of most aquatic crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s are biomineralized with calcium carbonate
Calcium carbonate
Calcium carbonate is a chemical compound with the formula CaCO3. It is a common substance found in rocks in all parts of the world, and is the main component of shells of marine organisms, snails, coal balls, pearls, and eggshells. Calcium carbonate is the active ingredient in agricultural lime,...
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. Biomineralization generally affects the exocuticle and the outer part of the endocuticle. Two recent hypotheses about the evolution of biomineralization in arthropods and other groups of animals propose that it provides tougher defensive armor, and that it allows animals to grow larger and stronger by providing more rigid skeletons; and in either case a mineral-organic composite
Composite material
Composite materials, often shortened to composites or called composition materials, are engineered or naturally occurring materials made from two or more constituent materials with significantly different physical or chemical properties which remain separate and distinct at the macroscopic or...
exoskeleton is cheaper to build than an all-organic one of comparable strength.
The cuticle can have seta
Seta
Seta is a biological term derived from the Latin word for "bristle". It refers to a number of different bristle- or hair-like structures on living organisms.-Animal setae:In zoology, most "setae" occur in invertebrates....
e (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
Feather
Feathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds and some non-avian theropod dinosaurs. They are considered the most complex integumentary structures found in vertebrates, and indeed a premier example of a complex evolutionary novelty. They...
-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
Felt
Felt is a non-woven cloth that is produced by matting, condensing and pressing woollen fibres. While some types of felt are very soft, some are tough enough to form construction materials. Felt can be of any colour, and made into any shape or size....
-like coats of setae to trap air, extending the time they can spend under water; heavy, rigid setae serve as defensive spines.
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; for example, all spiders extend their legs hydraulically and can generate pressures up to eight times their resting level.
Molting
The exoskeleton cannot stretch and thus restricts growth. Arthropods therefore replace their exoskeletons by molting, or shedding the old exoskeleton after growing a new one that is not yet hardened. Molting cycles run nearly continuously until an arthropod reaches full size.In the initial phase of molting, the animal stops feeding and its epidermis releases molting fluid, a mixture of enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
s that digests the endocuticle and thus detaches the old cuticle. This phase begins when the epidermis
Epidermis (zoology)
The Epidermis is an epithelium that covers the body of an eumetazoan . Eumetazoa have a cavity lined with a similar epithelium, the gastrodermis, which forms a boundary with the epidermis at the mouth.Sponges have no epithelium, and therefore no epidermis or gastrodermis...
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.
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. Molting may be responsible for 80 to 90% of all arthropod deaths.
Internal organs
Arthropod bodies are also segmented internally, and the nervous, muscular, circulatory, and excretory systems have repeated components. Arthropods come from a lineage of animals that have a coelomCoelom
The coelom is a fluid-filled cavity formed within the mesoderm. Coeloms developed in triploblasts but were subsequently lost in several lineages. Loss of coelom is correlated with reduction in body size...
, 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
Hydrostatic skeleton
A hydrostatic skeleton or hydroskeleton is a structure found in many cold-blooded organisms and soft-bodied animals consisting of a fluid-filled cavity, the coelom, surrounded by muscles. The pressure of the fluid and action of the surrounding circular and longitudinal muscles are used to change an...
, 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
Blood
Blood is a specialized bodily fluid in animals that delivers necessary substances such as nutrients and oxygen to the cells and transports metabolic waste products away from those same cells....
flows.
Arthropods have open circulatory system
Circulatory system
The circulatory system is an organ system that passes nutrients , gases, hormones, blood cells, etc...
s, although most have a few short, open-ended arteries
Artery
Arteries are blood vessels that carry blood away from the heart. This blood is normally oxygenated, exceptions made for the pulmonary and umbilical arteries....
. In chelicerates and crustaceans, the blood carries oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...
to the tissues, while hexapods
Hexapoda
The subphylum Hexapoda constitutes the largest grouping of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura . The Collembola are very abundant in terrestrial environments...
use a separate system of tracheae
Invertebrate trachea
The invertebrate trachea refers to the open respiratory system composed of spiracles, tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues....
. Many crustaceans, but few chelicerates and tracheates, use respiratory pigment
Respiratory pigment
A respiratory pigment is a molecule, such as hemoglobin in humans, that increases the oxygen-carrying capacity of the blood. The four most common invertebrate respiratory pigments are hemoglobin, haemocyanin, haemerythrin and chlorocruorin...
s to assist oxygen transport. The most common respiratory pigment in arthropods is copper
Copper
Copper is a chemical element with the symbol Cu and atomic number 29. It is a ductile metal with very high thermal and electrical conductivity. Pure copper is soft and malleable; an exposed surface has a reddish-orange tarnish...
-based hemocyanin
Hemocyanin
Hemocyanins are respiratory proteins in the form of metalloproteins containing two copper atoms that reversibly bind a single oxygen molecule . Oxygenation causes a color change between the colorless Cu deoxygenated form and the blue Cu oxygenated form...
; this is used by many crustaceans and a few centipede
Centipede
Centipedes are arthropods belonging to the class Chilopoda of the subphylum Myriapoda. They are elongated metameric animals with one pair of legs per body segment. Despite the name, centipedes can have a varying number of legs from under 20 to over 300. Centipedes have an odd number of pairs of...
s. A few crustaceans and insects use iron-based hemoglobin
Hemoglobin
Hemoglobin is the iron-containing oxygen-transport metalloprotein in the red blood cells of all vertebrates, with the exception of the fish family Channichthyidae, as well as the tissues of some invertebrates...
, the respiratory pigment used by vertebrate
Vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...
s. As with other invertebrates and unlike among vertebrates, the respiratory pigments of those arthropods that have them are generally dissolved in the blood and rarely enclosed in corpuscles
Blood cell
A blood cell, also called a hematocyte, is a cell normally found in blood. In mammals, these fall into three general categories:* red blood cells — Erythrocytes* white blood cells — Leukocytes* platelets — Thrombocytes...
.
The heart is typically 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 ligament
Ligament
In anatomy, the term ligament is used to denote any of three types of structures. Most commonly, it refers to fibrous tissue that connects bones to other bones and is also known as articular ligament, articular larua, fibrous ligament, or true ligament.Ligament can also refer to:* Peritoneal...
s or by small muscle
Muscle
Muscle is a contractile tissue of animals and is derived from the mesodermal layer of embryonic germ cells. Muscle cells contain contractile filaments that move past each other and change the size of the cell. They are classified as skeletal, cardiac, or smooth muscles. Their function is to...
s, 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.
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 lung
Book lung
A book lung is a type of respiration organ used for atmospheric gas exchange and is found in arachnids, such as scorpions and spiders. Each of these organs is found inside a ventral abdominal cavity and connects with the surroundings through a small opening. Book lungs are not related to the lungs...
s. 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 arachnid
Arachnid
Arachnids are a class of joint-legged invertebrate animals in the subphylum Chelicerata. All arachnids have eight legs, although in some species the front pair may convert to a sensory function. The term is derived from the Greek words , meaning "spider".Almost all extant arachnids are terrestrial...
s.
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
Sensory nerve
Sensory nerves are nerves that receive sensory stimuli, such as how something feels and if it is painful, smooth, rough, etc.They are made up of nerve fibers, called sensory fibers .Sensory neurons are neurons that are activated by sensory input Sensory nerves are nerves that receive 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 commissure
Commissure
A commissure is the place where two things are joined. The term is used especially in the fields of anatomy and biology.In anatomy, commissure refers to a bundle of nerve fibers that cross the midline at their level of origin or entry .* The most common usage of the term refers to the brain's...
s (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
Esophagus
The esophagus is an organ in vertebrates which consists of a muscular tube through which food passes from the pharynx to the stomach. During swallowing, food passes from the mouth through the pharynx into the esophagus and travels via peristalsis to the stomach...
. 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").
There are two different types of arthropod excretory systems. In aquatic arthropods, the end-product of biochemical reactions that metabolise
Metabolism
Metabolism is the set of chemical reactions that happen in the cells of living organisms to sustain life. These processes allow organisms to grow and reproduce, maintain their structures, and respond to their environments. Metabolism is usually divided into two categories...
nitrogen
Nitrogen
Nitrogen is a chemical element that has the symbol N, atomic number of 7 and atomic mass 14.00674 u. Elemental nitrogen is a colorless, odorless, tasteless, and mostly inert diatomic gas at standard conditions, constituting 78.08% by volume of Earth's atmosphere...
is ammonia
Ammonia
Ammonia is a compound of nitrogen and hydrogen with the formula . It is a colourless gas with a characteristic pungent odour. Ammonia contributes significantly to the nutritional needs of terrestrial organisms by serving as a precursor to food and fertilizers. Ammonia, either directly or...
, 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. 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. Various groups of terrestrial arthropods have independently developed a different system: the end-product of nitrogen metabolism is uric acid
Uric acid
Uric acid is a heterocyclic compound of carbon, nitrogen, oxygen, and hydrogen with the formula C5H4N4O3. It forms ions and salts known as urates and acid urates such as ammonium acid urate. Uric acid is created when the body breaks down purine nucleotides. High blood concentrations of uric acid...
, which can be excreted as dry material; the Malpighian tubule system
Malpighian tubule system
The Malpighian tubule system is a type of excretory and osmoregulatory system found in some Uniramia , arachnids and tardigrades....
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
Feces
Feces, faeces, or fæces is a waste product from an animal's digestive tract expelled through the anus or cloaca during defecation.-Etymology:...
. Most aquatic arthropods and some terrestrial ones also have organs called nephridia ("little kidney
Kidney
The kidneys, organs with several functions, serve essential regulatory roles in most animals, including vertebrates and some invertebrates. They are essential in the urinary system and also serve homeostatic functions such as the regulation of electrolytes, maintenance of acid–base balance, and...
s"), which extract other wastes for excretion as urine
Urine
Urine is a typically sterile liquid by-product of the body that is secreted by the kidneys through a process called urination and excreted through the urethra. Cellular metabolism generates numerous by-products, many rich in nitrogen, that require elimination from the bloodstream...
.
Senses
The stiff cuticleCuticle
A cuticle , or cuticula, is a term used for any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticles" are non-homologous; differing in their origin, structure, function, and chemical composition...
s 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 seta
Seta
Seta is a biological term derived from the Latin word for "bristle". It refers to a number of different bristle- or hair-like structures on living organisms.-Animal setae:In zoology, most "setae" occur in invertebrates....
e, respond to different levels of force, from strong contact to very weak air currents. Chemical sensors provide equivalents of taste
Taste
Taste is one of the traditional five senses. It refers to the ability to detect the flavor of substances such as food, certain minerals, and poisons, etc....
and smell
Olfaction
Olfaction is the sense of smell. This sense is mediated by specialized sensory cells of the nasal cavity of vertebrates, and, by analogy, sensory cells of the antennae of invertebrates...
, often by means of setae. Pressure sensors often take the form of membranes that function as eardrum
Eardrum
The eardrum, or tympanic membrane, is a thin membrane that separates the external ear from the middle ear in humans and other tetrapods. Its function is to transmit sound from the air to the ossicles inside the middle ear. The malleus bone bridges the gap between the eardrum and the other ossicles...
s, but are connected directly to nerves rather than to auditory ossicles. The antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
of most hexapods include sensor packages that monitor humidity
Humidity
Humidity is a term for the amount of water vapor in the air, and can refer to any one of several measurements of humidity. Formally, humid air is not "moist air" but a mixture of water vapor and other constituents of air, and humidity is defined in terms of the water content of this mixture,...
, moisture and temperature.
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s are pigment-cup ocelli that are capable of forming images, and those of jumping spider
Jumping spider
The jumping spider family contains more than 500 described genera and about 5,000 described species, making it the largest family of spiders with about 13% of all species. Jumping spiders have some of the best vision among invertebrates and use it in courtship, hunting and navigation...
s can rotate to track prey.
Compound eyes consist of fifteen to several thousand independent ommatidia, columns that are usually hexagonal in cross section
Cross section (geometry)
In geometry, a cross-section is the intersection of a figure in 2-dimensional space with a line, or of a body in 3-dimensional space with a plane, etc...
. Each ommatidium is an independent sensor, with its own light-sensitive cells and often with its own lens
Lens (anatomy)
The crystalline lens is a transparent, biconvex structure in the eye that, along with the cornea, helps to refract light to be focused on the retina. The lens, by changing shape, functions to change the focal distance of the eye so that it can focus on objects at various distances, thus allowing a...
and cornea
Cornea
The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Together with the lens, the cornea refracts light, with the cornea accounting for approximately two-thirds of the eye's total optical power. In humans, the refractive power of the cornea is...
. Compound eyes have a wide field of view, and can detect fast movement and, in some cases, the polarization of light. 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 centimetres (7.9 in) are most important to most arthropods. 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.
Most arthropods lack balance and acceleration
Acceleration
In physics, acceleration is the rate of change of velocity with time. In one dimension, acceleration is the rate at which something speeds up or slows down. However, since velocity is a vector, acceleration describes the rate of change of both the magnitude and the direction of velocity. ...
sensors, and rely on their eyes to tell them which way is up. The self-righting behavior of cockroach
Cockroach
Cockroaches are insects of the order Blattaria or Blattodea, of which about 30 species out of 4,500 total are associated with human habitations...
es is triggered when pressure sensors on the underside of the feet report no pressure. However many malacostracan crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s have statocyst
Statocyst
The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, echinoderms, cephalopods, and crustaceans. A similar structure is also found in Xenoturbella. The statocyst consists of a sac-like structure containing a mineralised mass and numerous...
s, which provide the same sort of information as the balance and motion sensors of the vertebrate inner ear
Inner ear
The inner ear is the innermost part of the vertebrate ear. In mammals, it consists of the bony labyrinth, a hollow cavity in the temporal bone of the skull with a system of passages comprising two main functional parts:...
.
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.
Reproduction and development
Barnacle
A barnacle is a type of arthropod belonging to infraclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in erosive settings. They are sessile suspension feeders, and have...
s, are hermaphroditic, that is, each can have the organs of both sex
Sex
In biology, sex is a process of combining and mixing genetic traits, often resulting in the specialization of organisms into a male or female variety . Sexual reproduction involves combining specialized cells to form offspring that inherit traits from both parents...
es. However, individuals of most species remain of one sex all their lives. A few species of insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s and crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s can reproduce by parthenogenesis
Parthenogenesis
Parthenogenesis is a form of asexual reproduction found in females, where growth and development of embryos occur without fertilization by a male...
, for example, without mating, especially if conditions favor a "population explosion". However most arthropods rely on sexual reproduction
Sexual reproduction
Sexual reproduction is the creation of a new organism by combining the genetic material of two organisms. There are two main processes during sexual reproduction; they are: meiosis, involving the halving of the number of chromosomes; and fertilization, involving the fusion of two gametes and the...
, and parthenogenetic species often revert to sexual reproduction when conditions become less favorable. Aquatic
Aquatic animal
An aquatic animal is an animal, either vertebrate or invertebrate, which lives in water for most or all of its life. It may breathe air or extract its oxygen from that dissolved in water through specialised organs called gills, or directly through its skin. Natural environments and the animals that...
arthropods may breed by external fertilization
External fertilization
External fertilization is a form of fertilization in which a sperm cell is united with an egg cell external to the bodies of the reproducing individuals. In contrast, internal fertilization takes place inside the female after insemination through copulation....
, as for example frog
Frog
Frogs are amphibians in the order Anura , formerly referred to as Salientia . Most frogs are characterized by a short body, webbed digits , protruding eyes and the absence of a tail...
s also do, or by internal fertilization
Internal fertilization
In mammals, internal fertilization is done through copulation, which involves the insertion of the penis into the vagina. Some other higher vertebrate animals reproduce internally, but their fertilization is cloacal.The union of spermatozoa of the parent organism. At some point, the growing egg or...
, where the ova
Ovum
An ovum is a haploid female reproductive cell or gamete. Both animals and embryophytes have ova. The term ovule is used for the young ovum of an animal, as well as the plant structure that carries the female gametophyte and egg cell and develops into a seed after fertilization...
remain in the female's body and the sperm
Sperm
The term sperm is derived from the Greek word sperma and refers to the male reproductive cells. In the types of sexual reproduction known as anisogamy and oogamy, there is a marked difference in the size of the gametes with the smaller one being termed the "male" or sperm cell...
must somehow be inserted. All known terrestrial arthropods use internal fertilization, as unprotected sperm
Sperm
The term sperm is derived from the Greek word sperma and refers to the male reproductive cells. In the types of sexual reproduction known as anisogamy and oogamy, there is a marked difference in the size of the gametes with the smaller one being termed the "male" or sperm cell...
and ova would not survive long in these environments. In a few cases the sperm transfer is direct from the male's penis
Penis
The penis is a biological feature of male animals including both vertebrates and invertebrates...
to the female's oviduct
Oviduct
In non-mammalian vertebrates, the passageway from the ovaries to the outside of the body is known as the oviduct. The eggs travel along the oviduct. These eggs will either be fertilized by sperm to become a zygote, or will degenerate in the body...
, but it is more often indirect. Some crustaceans and spiders use modified appendages to transfer the sperm to the female. On the other hand, many male terrestrial
Terrestrial
Terrestrial refers to things related to land or the planet Earth.Terrestrial may also refer to:* Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to in trees etc.** A fishing fly that simulates the...
arthropods produce spermatophore
Spermatophore
A spermatophore or sperm ampulla is a capsule or mass created by males of various animal species, containing spermatozoa and transferred in entirety to the female's ovipore during copulation...
s, 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
Courtship
Courtship is the period in a couple's relationship which precedes their engagement and marriage, or establishment of an agreed relationship of a more enduring kind. In courtship, a couple get to know each other and decide if there will be an engagement or other such agreement...
rituals look likely to be successful.
Silverfish
Lepisma saccharina, frequently called silverfish, fishmoths, carpet sharks or paramites, are small, wingless insects in the order Thysanura...
, the hatchlings do not feed and may be helpless until after their first molt. Many insects hatch as grubs or caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...
s, which do not have segmented limbs or hardened cuticles, and metamorphose
Metamorphosis
Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation...
into adult forms by entering an inactive phase in which the larval tissues are broken down and re-used to build the adult body. Dragonfly
Dragonfly
A dragonfly is a winged insect belonging to the order Odonata, the suborder Epiprocta or, in the strict sense, the infraorder Anisoptera . It is characterized by large multifaceted eyes, two pairs of strong transparent wings, and an elongated body...
larvae have the typical cuticles and jointed limbs of arthropods but are flightless water-breathers with extendable jaws. Crustaceans commonly hatch as tiny nauplius larvae that have only three segments and pairs of appendages.
Last common ancestor
The last common ancestor of all arthropods is reconstructed as a modular organism with each module covered by its own scleriteSclerite
A sclerite is a hardened body part. The term is used in various branches of biology for various structures including hardened portions of sponges, but it is most commonly used for the hardened portions of arthropod exoskeletons....
(armor plate) and bearing a pair of biramous limbs. Whether the ancestral limb was uniramous or biramous is far from a settled debate, though.
This Ur-arthropod had a ventral mouth, pre-oral antennae and dorsal eyes at the front of the body. It was a non-discriminatory sediment
Sediment
Sediment is naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of fluids such as wind, water, or ice, and/or by the force of gravity acting on the particle itself....
feeder, processing whatever sediment came its way for food.
Fossil record
Ediacaran
The Ediacaran Period , named after the Ediacara Hills of South Australia, is the last geological period of the Neoproterozoic Era and of the Proterozoic Eon, immediately preceding the Cambrian Period, the first period of the Paleozoic Era and of the Phanerozoic Eon...
animals Parvancorina
Parvancorina
Parvancorina is a genus of shield-shaped Ediacaran fossils. It has a raised ridge down the central axis of symmetry. This ridge can be high in unflattened fossils. At the 'head' end of the ridge there are two quarter circle shaped raised arcs attached. In front of this are two nested...
and Spriggina
Spriggina
Fossils of Spriggina are known from the Ediacaran period, around . The segmented organism reached about 3 cm in length and may have been predatory...
, from around 555 Mya, were arthropods. Small arthropods with bivalve-like shells have been found in Early Cambrian fossil beds dating in China. The earliest Cambrian trilobite
Trilobite
Trilobites are a well-known fossil group of extinct marine arthropods that form the class Trilobita. The first appearance of trilobites in the fossil record defines the base of the Atdabanian stage of the Early Cambrian period , and they flourished throughout the lower Paleozoic era before...
fossils are about 530 million years old, but the class was already quite diverse and worldwide, suggesting that they had been around for quite some time. Re-examination in the 1970s of the Burgess Shale
Burgess Shale
The Burgess Shale Formation, located in the Canadian Rockies of British Columbia, is one of the world's most celebrated fossil fields, and the best of its kind. It is famous for the exceptional preservation of the soft parts of its fossils...
fossils from about 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
Cambrian explosion
The Cambrian explosion or Cambrian radiation was the relatively rapid appearance, around , of most major phyla, as demonstrated in the fossil record, accompanied by major diversification of other organisms, including animals, phytoplankton, and calcimicrobes...
. A fossil of Marrella from the Burgess Shale has provided the earliest clear evidence of molting.
The earliest fossil crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s date from about in the Cambrian
Cambrian
The Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...
, and fossil shrimp
Shrimp
Shrimp are swimming, decapod crustaceans classified in the infraorder Caridea, found widely around the world in both fresh and salt water. Adult shrimp are filter feeding benthic animals living close to the bottom. They can live in schools and can swim rapidly backwards. Shrimp are an important...
from about apparently formed a tight-knit procession across the seabed. Crustacean fossils are common from the Ordovician
Ordovician
The Ordovician is a geologic period and system, the second of six of the Paleozoic Era, and covers the time between 488.3±1.7 to 443.7±1.5 million years ago . It follows the Cambrian Period and is followed by the Silurian Period...
period onwards. They have remained almost entirely aquatic, possibly because they never developed excretory system
Excretory system
The excretory system is a passive biological system that removes excess, unnecessary or dangerous materials from an organism, so as to help maintain homeostasis within the organism and prevent damage to the body. It is responsible for the elimination of the waste products of metabolism as well as...
s that conserve water.
Arthropods provide the earliest identifiable fossils of land animals, from about in the Late Silurian
Silurian
The Silurian is a geologic period and system that extends from the end of the Ordovician Period, about 443.7 ± 1.5 Mya , to the beginning of the Devonian Period, about 416.0 ± 2.8 Mya . As with other geologic periods, the rock beds that define the period's start and end are well identified, but the...
, and terrestrial tracks from about appear to have been made by arthropods. Arthropods were well pre-adapted
Preadaptation
In evolutionary biology, preadaptation describes a situation where a species evolves to use a preexisting structure or trait inherited from an ancestor for a potentially unrelated function...
to colonize land, because their existing jointed exoskeletons provided protection against desiccation, support against gravity and a means of locomotion that was not dependent on water. Around the same time the aquatic, scorpion-like eurypterid
Eurypterid
Eurypterids are an extinct group of arthropods related to arachnids which include the largest known arthropods that ever lived. They are members of the extinct order Eurypterida ; which is the most diverse Paleozoic chelicerate order in terms of species...
s became the largest ever arthropods, some as long as 2.5 metres (8.2 ft).
The oldest known arachnid
Arachnid
Arachnids are a class of joint-legged invertebrate animals in the subphylum Chelicerata. All arachnids have eight legs, although in some species the front pair may convert to a sensory function. The term is derived from the Greek words , meaning "spider".Almost all extant arachnids are terrestrial...
is the trigonotarbid Palaeotarbus jerami, from about in the Silurian
Silurian
The Silurian is a geologic period and system that extends from the end of the Ordovician Period, about 443.7 ± 1.5 Mya , to the beginning of the Devonian Period, about 416.0 ± 2.8 Mya . As with other geologic periods, the rock beds that define the period's start and end are well identified, but the...
period. Attercopus
Attercopus
Previously interpreted as the world's oldest spider, Attercopus fimbriunguis belongs to an extinct order of arachnids named Uraraneida; thought to be close to the origins of spiders...
fimbriunguis, from in the Devonian
Devonian
The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...
period, bears the earliest known silk-producing spigots, but its lack of spinnerets means it was not one of the true spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s, which first appear in the Late Carboniferous
Carboniferous
The Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...
over . The Jurassic
Jurassic
The Jurassic is a geologic period and system that extends from about Mya to Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...
and Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
periods provide a large number of fossil spiders, including representatives of many modern families. Fossils of aquatic scorpion
Scorpion
Scorpions are predatory arthropod animals of the order Scorpiones within the class Arachnida. They have eight legs and are easily recognized by the pair of grasping claws and the narrow, segmented tail, often carried in a characteristic forward curve over the back, ending with a venomous stinger...
s with gill
Gill
A gill is a respiratory organ found in many aquatic organisms that extracts dissolved oxygen from water, afterward excreting carbon dioxide. The gills of some species such as hermit crabs have adapted to allow respiration on land provided they are kept moist...
s appear in the Silurian and Devonian
Devonian
The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...
periods, and the earliest fossil of an air-breathing scorpion with book lung
Book lung
A book lung is a type of respiration organ used for atmospheric gas exchange and is found in arachnids, such as scorpions and spiders. Each of these organs is found inside a ventral abdominal cavity and connects with the surroundings through a small opening. Book lungs are not related to the lungs...
s dates from the Early Carboniferous period.
The oldest definitive insect fossil is the Devonian
Devonian
The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...
Rhyniognatha hirsti, dated at , but its mandibles are of a type found only in winged insects, which suggests that the earliest insects appeared in the Silurian period. The Mazon Creek lagerstätten
Mazon Creek fossils
The Mazon Creek fossil beds are a conservation found near Morris, in Grundy County, Illinois. The fossil beds are located in ironstone concretions, formed approximately in the mid-Pennsylvanian epoch of the Carboniferous period...
from the Late Carboniferous, about , include about 200 species, some gigantic by modern standards, and indicate that insects had occupied their main modern ecological niche
Ecological niche
In ecology, a niche is a term describing the relational position of a species or population in its ecosystem to each other; e.g. a dolphin could potentially be in another ecological niche from one that travels in a different pod if the members of these pods utilize significantly different food...
s as herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...
s, detritivore
Detritivore
Detritivores, also known as detritophages or detritus feeders or detritus eaters or saprophages, are heterotrophs that obtain nutrients by consuming detritus . By doing so, they contribute to decomposition and the nutrient cycles...
s and insectivore
Insectivore
An insectivore is a type of carnivore with a diet that consists chiefly of insects and similar small creatures. An alternate term is entomophage, which also refers to the human practice of eating insects....
s. Social termites and ants first appear in the Early Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
, and advanced social bees have been found in Late Cretaceous rocks but did not become abundant until the Mid Cenozoic
Cenozoic
The Cenozoic era is the current and most recent of the three Phanerozoic geological eras and covers the period from 65.5 mya to the present. The era began in the wake of the Cretaceous–Tertiary extinction event at the end of the Cretaceous that saw the demise of the last non-avian dinosaurs and...
.
Evolutionary family tree
Sidnie Manton
Sidnie Milana Manton, FRS was a British entomologist.-Early life:Sidnie Milana Manton was born in Kensington, London the daughter of a dentist. Her sister was Irene Manton , a botanist, She was educated in Kensington and at St...
and others argued that arthropods are polyphyletic, in other words, 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 spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s and scorpion
Scorpion
Scorpions are predatory arthropod animals of the order Scorpiones within the class Arachnida. They have eight legs and are easily recognized by the pair of grasping claws and the narrow, segmented tail, often carried in a characteristic forward curve over the back, ending with a venomous stinger...
s; the crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s; and the uniramia
Uniramia
S. M. Manton's polyphyletic Arthropod hypothesis with Uniramia as one of three Phyla.Uniramia is a group within the arthropods. In the past this group included the Onychophora, which are now considered a separate category. The group is currently used in a narrower sense.Uniramia is one of three...
, consisting of onychophorans, myriapods and hexapods
Hexapoda
The subphylum Hexapoda constitutes the largest grouping of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura . The Collembola are very abundant in terrestrial environments...
. These arguments usually bypassed trilobite
Trilobite
Trilobites are a well-known fossil group of extinct marine arthropods that form the class Trilobita. The first appearance of trilobites in the fossil record defines the base of the Atdabanian stage of the Early Cambrian period , and they flourished throughout the lower Paleozoic era before...
s, 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
Convergent evolution
Convergent evolution describes the acquisition of the same biological trait in unrelated lineages.The wing is a classic example of convergent evolution in action. Although their last common ancestor did not have wings, both birds and bats do, and are capable of powered flight. The wings are...
, as natural consequences of having rigid, segmented exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
s; 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.
including Hallucigenia
Hallucigenia
Hallucigenia is an extinct genus of animal found as fossils in the Middle Cambrian-aged Burgess Shale formation of British Columbia, Canada, represented by the species H. sparsa, and in the Lower Cambrian Maotianshan shale of China, represented by the species H. fortis...
and Microdictyon
Microdictyon
Microdictyon is an extinct "armored worm" coated with dot-likescleritic scales, known from the Early CambrianMaotianshan shale of Yunnan China. Microdictyon is sometimes included...
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Further analysis and discoveries in the 1990s reversed this view, and led to acceptance that arthropods are monophyletic, in other words they do share a common ancestor that was itself an arthropod. For example Graham Budd
Graham Budd
Graham Edward Budd is a British palaeontologist, Professor of paleobiology at Uppsala University.Budd’s research primarily has focused on the anatomy and evolutionary significance of Paleozoic arthropods and in the integration of palaeontology into evolutionary developmental biology...
's analyses of Kerygmachela in 1993 and of Opabinia
Opabinia
Opabinia is an animal genus found in Cambrian fossil deposits. Its sole species, Opabinia regalis, is known from the Middle Cambrian Burgess Shale of British Columbia, Canada. Fewer than twenty good specimens have been described; 3 specimens of Opabinia are known from the Greater Phyllopod bed,...
in 1996 convinced him that these animals were similar to onychophorans and to various Early Cambrian
Cambrian
The Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...
"lobopods", and he presented an "evolutionary family tree" that showed these as "aunts" and "cousins" of all arthropods. These changes made the scope of the term "arthropod" unclear, and Claus Nielsen proposed that the wider group should be labelled "Panarthropoda
Panarthropoda
Panarthropoda is a taxon combining the phyla Arthropoda, Tardigrada, and Onychophora. Dunn et al. in 2008 suggested that the tardigrades could be grouped along with the nematodes, leaving Onychophora as the sister group to the arthropods...
" ("all the arthropods") while the animals with jointed limbs and hardened cuticles should be called "Euarthropoda" ("true arthropods").
A contrary view was presented in 2003, when Jan Bergström and Xian-Guang Hou 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. This reasoning implies that Parapeytoia
Parapeytoia
Parapeytoia was a prehistoric animal that lived over 530 million years ago that lived in Maotianshan shales of prehistoric China. Like the anomalocarids that it resembled , Parapeytoia had two head appendages in front of the also anomalocarid like round mouth, eyes on stalks, and fleshy lobed...
, which has legs and a backward-pointing mouth like that of the earliest arthropods, is a more credible closest relative of arthropods than is Anomalocaris
Anomalocaris
Anomalocaris is an extinct genus of anomalocaridid, which are, in turn, thought to be closely related to the arthropods. The first fossils of Anomalocaris were discovered in the Ogygopsis Shale by Joseph Frederick Whiteaves, with more examples found by Charles Doolittle Walcott in the famed...
. In 2006, they suggested that arthropods were more closely related to lobopods and tardigrade
Tardigrade
Tardigrades form the phylum Tardigrada, part of the superphylum Ecdysozoa. They are microscopic, water-dwelling, segmented animals with eight legs. Tardigrades were first described by Johann August Ephraim Goeze in 1773...
s than to anomalocarids.
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
Articulata Hypothesis
The Articulata Hypothesis is the grouping in a higher taxon of animals with segmented bodies, consisting of Annelida and Panarthropoda. This theory states that these groups are descended from a common segmented ancestor...
. There had been competing proposals that arthropods were closely related to other groups such as nematode
Nematode
The nematodes or roundworms are the most diverse phylum of pseudocoelomates, and one of the most diverse of all animals. Nematode species are very difficult to distinguish; over 28,000 have been described, of which over 16,000 are parasitic. It has been estimated that the total number of nematode...
s, priapulids and tardigrade
Tardigrade
Tardigrades form the phylum Tardigrada, part of the superphylum Ecdysozoa. They are microscopic, water-dwelling, segmented animals with eight legs. Tardigrades were first described by Johann August Ephraim Goeze in 1773...
s, 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
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...
sequences produced a coherent scheme showing arthropods as members of a superphylum labelled Ecdysozoa
Ecdysozoa
Ecdysozoa is a group of protostome animals, including Arthropoda , Nematoda, and several smaller phyla. They were first defined by Aguinaldo et al. in 1997, based mainly on trees constructed using 18S ribosomal RNA genes...
("animals that molt"), 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 brachiopod
Brachiopod
Brachiopods are a phylum of marine animals that have hard "valves" on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection...
s in another superphylum, Lophotrochozoa
Lophotrochozoa
The Lophotrochozoa are a major grouping of protostome animals. The taxon was discovered based on molecular data. Molecular evidence such as a result of studies of the evolution of small-subunit ribosomal RNA supports the monophyly of the phyla listed in the infobox shown at right.-Terminology:The...
.
If the Ecdysozoa hypothesis is correct, then segmentation of arthropods and annelids either has evolved convergently
Convergent evolution
Convergent evolution describes the acquisition of the same biological trait in unrelated lineages.The wing is a classic example of convergent evolution in action. Although their last common ancestor did not have wings, both birds and bats do, and are capable of powered flight. The wings are...
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.
Classification
Euarthropods are typically classified into five subphylaSubphylum
In life, a subphylum is a taxonomic rank intermediate between phylum and superclass. The rank of subdivision in plants and fungi is equivalent to subphylum.Not all phyla are divided into subphyla...
, of which one is extinct:
- TrilobiteTrilobiteTrilobites are a well-known fossil group of extinct marine arthropods that form the class Trilobita. The first appearance of trilobites in the fossil record defines the base of the Atdabanian stage of the Early Cambrian period , and they flourished throughout the lower Paleozoic era before...
s are a group of formerly numerous marine animals that disappeared in the Permian-Triassic extinction eventPermian-Triassic extinction eventThe Permian–Triassic extinction event, informally known as the Great Dying, was an extinction event that occurred 252.28 Ma ago, forming the boundary between the Permian and Triassic geologic periods, as well as the Paleozoic and Mesozoic eras...
, though they were in decline prior to this killing blow, having been reduced to one order in the Late Devonian extinctionLate Devonian extinctionThe Late Devonian extinction was one of five major extinction events in the history of the Earth's biota. A major extinction, the Kellwasser Event, occurred at the boundary that marks the beginning of the last phase of the Devonian period, the Famennian faunal stage, , about 374 million years ago...
. - CheliceratesChelicerataThe subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda, and includes horseshoe crabs, scorpions, spiders and mites...
include spiderSpiderSpiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s, miteMiteMites, along with ticks, are small arthropods belonging to the subclass Acari and the class Arachnida. The scientific discipline devoted to the study of ticks and mites is called acarology.-Diversity and systematics:...
s, scorpionScorpionScorpions are predatory arthropod animals of the order Scorpiones within the class Arachnida. They have eight legs and are easily recognized by the pair of grasping claws and the narrow, segmented tail, often carried in a characteristic forward curve over the back, ending with a venomous stinger...
s and related organisms. They are characterised by the presence of cheliceraeCheliceraeThe chelicerae are mouthparts of the Chelicerata, an arthropod subphylum that includes arachnids, Merostomata , and Pycnogonida . Chelicerae are pointed appendages which are used to grasp food, and are found in place of the chewing mandibles most other arthropods have...
, appendages just above / in front of the mouth. Chelicerae appear in scorpions as tiny claws that they use in feeding, but those of spiders have developed as fangs that inject venom. - MyriapodsMyriapodaMyriapoda is a subphylum of arthropods containing millipedes, centipedes, and others. The group contains 13,000 species, all of which are terrestrial...
comprise millipedeMillipedeMillipedes are arthropods that have two pairs of legs per segment . Each segment that has two pairs of legs is a result of two single segments fused together as one...
s, centipedeCentipedeCentipedes are arthropods belonging to the class Chilopoda of the subphylum Myriapoda. They are elongated metameric animals with one pair of legs per body segment. Despite the name, centipedes can have a varying number of legs from under 20 to over 300. Centipedes have an odd number of pairs of...
s, and their relatives and have many body segments, each bearing one or two pairs of legs. They are sometimes grouped with the hexapods. - HexapodsHexapodaThe subphylum Hexapoda constitutes the largest grouping of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura . The Collembola are very abundant in terrestrial environments...
comprise insectInsectInsects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s and three small orders of insect-like animals with six thoracic legs. They are sometimes grouped with the myriapods, in a group called UniramiaUniramiaS. M. Manton's polyphyletic Arthropod hypothesis with Uniramia as one of three Phyla.Uniramia is a group within the arthropods. In the past this group included the Onychophora, which are now considered a separate category. The group is currently used in a narrower sense.Uniramia is one of three...
, though genetic evidence tends to support a closer relationship between hexapods and crustaceans. - CrustaceanCrustaceanCrustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s are primarily aquatic (a notable exception being woodliceWoodlouseA woodlouse is a crustacean with a rigid, segmented, long exoskeleton and fourteen jointed limbs...
) and are characterised by having biramous appendages. They include lobsterLobsterClawed lobsters comprise a family of large marine crustaceans. Highly prized as seafood, lobsters are economically important, and are often one of the most profitable commodities in coastal areas they populate.Though several groups of crustaceans are known as lobsters, the clawed lobsters are most...
s, crabCrabTrue crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...
s, barnacleBarnacleA barnacle is a type of arthropod belonging to infraclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in erosive settings. They are sessile suspension feeders, and have...
s, crayfishCrayfishCrayfish, crawfish, or crawdads – members of the superfamilies Astacoidea and Parastacoidea – are freshwater crustaceans resembling small lobsters, to which they are related...
, shrimpShrimpShrimp are swimming, decapod crustaceans classified in the infraorder Caridea, found widely around the world in both fresh and salt water. Adult shrimp are filter feeding benthic animals living close to the bottom. They can live in schools and can swim rapidly backwards. Shrimp are an important...
and many others.
Aside from these major groups, there are also a number of fossil forms, mostly from the Early Cambrian
Cambrian
The Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...
, which are difficult to place, 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.
The phylogeny
Phylogenetics
In biology, phylogenetics is the study of evolutionary relatedness among groups of organisms , which is discovered through molecular sequencing data and morphological data matrices...
of the major extant arthropod groups has been an area of considerable interest and dispute. The most recent studies tend to suggest a paraphyletic
Paraphyly
A group of taxa is said to be paraphyletic if the group consists of all the descendants of a hypothetical closest common ancestor minus one or more monophyletic groups of descendants...
Crustacea with different hexapod groups nested within it. Myriapoda
Myriapoda
Myriapoda is a subphylum of arthropods containing millipedes, centipedes, and others. The group contains 13,000 species, all of which are terrestrial...
is grouped with Chelicerata
Chelicerata
The subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda, and includes horseshoe crabs, scorpions, spiders and mites...
in some recent studies (forming Myriochelata
Myriochelata
The Myriochelata or Paradoxopoda, is a proposed grouping of arthropods comprising the Myriapoda and Chelicerata...
), and with Pancrustacea
Pancrustacea
Pancrustacea is a clade, comprising all crustaceans and hexapods. This grouping is contrary to the Atelocerata hypothesis, in which Myriapoda and Hexapoda are sister taxa, and Crustacea are only more distantly related. As of 2010, the Pancrustacea taxon is considered well accepted. The clade has...
in other studies (forming Mandibulata
Mandibulata
The Mandibulata or mandibulates is a clade of arthropods that comprises the extant subphyla Myriapoda , Crustacea and Hexapoda . Mandibulata is currently believed to be the sister group of the clade Arachnomorpha, which comprises the rest of arthropods...
). The placement of the extinct trilobite
Trilobite
Trilobites are a well-known fossil group of extinct marine arthropods that form the class Trilobita. The first appearance of trilobites in the fossil record defines the base of the Atdabanian stage of the Early Cambrian period , and they flourished throughout the lower Paleozoic era before...
s is also a frequent subject of dispute.
Since the International Code of Zoological Nomenclature
International Code of Zoological Nomenclature
The International Code of Zoological Nomenclature is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals...
recognises no priority above the rank of family, many of the higher-level groups can be referred to by a variety of different names.
Interaction with humans
CrustaceanCrustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s such as crab
Crab
True crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...
s, lobster
Lobster
Clawed lobsters comprise a family of large marine crustaceans. Highly prized as seafood, lobsters are economically important, and are often one of the most profitable commodities in coastal areas they populate.Though several groups of crustaceans are known as lobsters, the clawed lobsters are most...
s, crayfish
Crayfish
Crayfish, crawfish, or crawdads – members of the superfamilies Astacoidea and Parastacoidea – are freshwater crustaceans resembling small lobsters, to which they are related...
, shrimp
Shrimp
Shrimp are swimming, decapod crustaceans classified in the infraorder Caridea, found widely around the world in both fresh and salt water. Adult shrimp are filter feeding benthic animals living close to the bottom. They can live in schools and can swim rapidly backwards. Shrimp are an important...
s and prawn
Prawn
Prawns are decapod crustaceans of the sub-order Dendrobranchiata. There are 540 extant species, in seven families, and a fossil record extending back to the Devonian...
s have long been part of human cuisine
Cuisine
Cuisine is a characteristic style of cooking practices and traditions, often associated with a specific culture. Cuisines are often named after the geographic areas or regions that they originate from...
, and are now farmed on a large commercial scale. Insects and their grubs are at least as nutritious as meat, and are eaten both raw and cooked in many non-European cultures. Cooked tarantula
Tarantula
Tarantulas comprise a group of often hairy and often very large arachnids belonging to the family Theraphosidae, of which approximately 900 species have been identified. Some members of the same Suborder may also be called "tarantulas" in the common parlance. This article will restrict itself to...
s are considered a delicacy in Cambodia
Cambodia
Cambodia , officially known as the Kingdom of Cambodia, is a country located in the southern portion of the Indochina Peninsula in Southeast Asia...
, and by the Piaroa
Piaroa
The Piaroa are an indigenous American ethnic group living along the banks of the Orinoco River and its tributaries in present day Venezuela, and in a few scattered locations elsewhere in Venezuela and in Colombia...
Indians of southern Venezuela, after the highly irritant hairs – the spider's main defense system – are removed. Humans also unintentionally eat arthropods in other foods, and food safety regulations lay down acceptable contamination levels for different kinds of food material. The intentional cultivation of arthropods and other small animals for human food, referred to as minilivestock, is now emerging in animal husbandry
Animal husbandry
Animal husbandry is the agricultural practice of breeding and raising livestock.- History :Animal husbandry has been practiced for thousands of years, since the first domestication of animals....
as an ecologically sound concept.
However, the greatest contribution of arthropods to human food supply is by pollination
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...
: a 2008 study examined the 100 crops that FAO
Fão
Fão is a town in Esposende Municipality in Portugal....
lists as grown for food, and estimated pollination's economic value as €153 billion, or 9.5% of the value of world agricultural production used for human food in 2005. Besides pollinating, bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...
s produce honey
Honey
Honey is a sweet food made by bees using nectar from flowers. The variety produced by honey bees is the one most commonly referred to and is the type of honey collected by beekeepers and consumed by humans...
, which is the basis of a rapidly growing industry and international trade.
The red dye cochineal
Cochineal
The cochineal is a scale insect in the suborder Sternorrhyncha, from which the crimson-colour dye carmine is derived. A primarily sessile parasite native to tropical and subtropical South America and Mexico, this insect lives on cacti from the genus Opuntia, feeding on plant moisture and...
, produced from a Central American species of insect, was economically important to the Aztec
Aztec
The Aztec people were certain ethnic groups of central Mexico, particularly those groups who spoke the Nahuatl language and who dominated large parts of Mesoamerica in the 14th, 15th and 16th centuries, a period referred to as the late post-classic period in Mesoamerican chronology.Aztec is the...
s and Mayans
Maya civilization
The Maya is a Mesoamerican civilization, noted for the only known fully developed written language of the pre-Columbian Americas, as well as for its art, architecture, and mathematical and astronomical systems. Initially established during the Pre-Classic period The Maya is a Mesoamerican...
, and while the region was under Spanish
Spain
Spain , officially the Kingdom of Spain languages]] under the European Charter for Regional or Minority Languages. In each of these, Spain's official name is as follows:;;;;;;), is a country and member state of the European Union located in southwestern Europe on the Iberian Peninsula...
control, becoming Mexico
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...
's second most-lucrative export; and it is now regaining some of the ground it lost to synthetic competitors. The blood of horseshoe crabs contains a clotting agent Limulus Amebocyte Lysate
Limulus Amebocyte Lysate
Limulus amebocyte lysate is an aqueous extract of blood cells from the horseshoe crab, Limulus polyphemus. LAL reacts with bacterial endotoxin or lipopolysaccharide , which is a membrane component of Gram negative bacteria...
which is now used to test that antibiotics and kidney machines are free of dangerous bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
, and to detect spinal meningitis and some cancer
Cancer
Cancer , known medically as a malignant neoplasm, is a large group of different diseases, all involving unregulated cell growth. In cancer, cells divide and grow uncontrollably, forming malignant tumors, and invade nearby parts of the body. The cancer may also spread to more distant parts of the...
s. Forensic entomology
Forensic entomology
Forensic entomology is the application and study of insect and other arthropod biology to criminal matters. It is primarily associated with death investigations; however, it may also be used to detect drugs and poisons, determine the location of an incident, and find the presence and time of the...
uses evidence provided by arthropods to establish the time and sometimes the place of death of a human, and in some cases the cause. Recently insects have also gained attention as potential sources of drugs and other medicinal substances.
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 robot
Robot
A robot is a mechanical or virtual intelligent agent that can perform tasks automatically or with guidance, typically by remote control. In practice a robot is usually an electro-mechanical machine that is guided by computer and electronic programming. Robots can be autonomous, semi-autonomous or...
ics. The redundancy provided by segments allows arthropods and biomimetic robots to move normally even with damaged or lost appendages.
| Disease | Insect | Cases per year | Deaths per year |
|---|---|---|---|
| Malaria Malaria Malaria is a mosquito-borne infectious disease of humans and other animals caused by eukaryotic protists of the genus Plasmodium. The disease results from the multiplication of Plasmodium parasites within red blood cells, causing symptoms that typically include fever and headache, in severe cases... |
Anopheles Anopheles Anopheles is a genus of mosquito. There are approximately 460 recognized species: while over 100 can transmit human malaria, only 30–40 commonly transmit parasites of the genus Plasmodium, which cause malaria in humans in endemic areas... mosquito |
267 M | 1 to 2 M |
| Yellow fever Yellow fever Yellow fever is an acute viral hemorrhagic disease. The virus is a 40 to 50 nm enveloped RNA virus with positive sense of the Flaviviridae family.... |
Aedes Aedes Aedes is a genus of mosquito originally found in tropical and subtropical zones, but now found on all continents excluding Antarctica. Some species have been spread by human activity. Aedes albopictus, a most invasive species was recently spread to the New World, including the U.S., by the used... mosquito |
4,432 | 1,177 |
| Filariasis Filariasis Filariasis is a parasitic disease and is considered an infectious tropical disease, that is caused by thread-like nematodes belonging to the superfamily Filarioidea, also known as "filariae".... |
Culex Culex Culex is a genus of mosquito, and is important in that several species serve as vectors of important diseases, such as West Nile virus, filariasis, Japanese encephalitis, St. Louis encephalitis and avian malaria.... 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 carried by blood-sucking insects. Other blood-sucking insects infect livestock
Livestock
Livestock refers to one or more domesticated animals raised in an agricultural setting to produce commodities such as food, fiber and labor. The term "livestock" as used in this article does not include poultry or farmed fish; however the inclusion of these, especially poultry, within the meaning...
with diseases that kill many animals and greatly reduce the usefulness of others. Tick
Tick
Ticks are small arachnids in the order Ixodida, along with mites, constitute the subclass Acarina. Ticks are ectoparasites , living by hematophagy on the blood of mammals, birds, and sometimes reptiles and amphibians...
s can cause tick paralysis
Tick paralysis
Tick paralysis is the only tick-borne disease that is not caused by an infectious organism. The illness is caused by a neurotoxin produced in the tick's salivary gland. After prolonged attachment, the engorged tick transmits the toxin to its host...
and several parasite-borne diseases in humans. A few of the closely related mite
Mite
Mites, along with ticks, are small arthropods belonging to the subclass Acari and the class Arachnida. The scientific discipline devoted to the study of ticks and mites is called acarology.-Diversity and systematics:...
s also infest humans, causing intense itching, and others cause allergic
Allergy
An Allergy is a hypersensitivity disorder of the immune system. Allergic reactions occur when a person's immune system reacts to normally harmless substances in the environment. A substance that causes a reaction is called an allergen. These reactions are acquired, predictable, and rapid...
diseases, including hay fever
Hay Fever
Hay Fever is a comic play written by Noël Coward in 1924 and first produced in 1925 with Marie Tempest as the first Judith Bliss. Laura Hope Crews played the role in New York...
, asthma
Asthma
Asthma is the common chronic inflammatory disease of the airways characterized by variable and recurring symptoms, reversible airflow obstruction, and bronchospasm. Symptoms include wheezing, coughing, chest tightness, and shortness of breath...
and eczema
Eczema
Eczema is a form of dermatitis, or inflammation of the epidermis . In England, an estimated 5.7 million or about one in every nine people have been diagnosed with the disease by a clinician at some point in their lives.The term eczema is broadly applied to a range of persistent skin conditions...
.
Many species of arthropods, principally insects but also mites, are agricultural and forest pests. The mite Varroa destructor
Varroa destructor
Varroa destructor is an external parasitic mite that attacks honey bees Apis cerana and Apis mellifera. The disease caused by the mites is called varroatosis....
has become the largest single problem faced by beekeeper
Beekeeper
A beekeeper is a person who keeps honey bees for the purposes of securing commodities such as honey, beeswax, pollen, royal jelly; pollinating fruits and vegetables; raising queens and bees for sale to other farmers; and/or for purposes satisfying natural scientific curiosity...
s worldwide. Efforts to control arthropod pests by large-scale use of pesticide
Pesticide
Pesticides are substances or mixture of substances intended for preventing, destroying, repelling or mitigating any pest.A pesticide may be a chemical unicycle, biological agent , antimicrobial, disinfectant or device used against any pest...
s have caused long term effects on human health and on biodiversity
Biodiversity
Biodiversity is the degree of variation of life forms within a given ecosystem, biome, or an entire planet. Biodiversity is a measure of the health of ecosystems. Biodiversity is in part a function of climate. In terrestrial habitats, tropical regions are typically rich whereas polar regions...
. Increasing arthropod resistance
Pesticide resistance
Pesticide resistance is the adaptation of pest population targeted by a pesticide resulting in decreased susceptibility to that chemical. In other words, pests develop a resistance to a chemical through natural selection: the most resistant organisms are the ones to survive and pass on their...
to pesticides has led to the development of integrated pest management
Integrated Pest Management
Integrated pest management is an ecological approach to agricultural pest control that integrates pesticides/herbicides into a management system incorporating a range of practices for economic control of a pest...
using a wide range of meaures including biological control. Predatory mites may be useful in controlling some mite pests.
External links
- Venomous Arthropods chapter in United States Environmental Protection AgencyUnited States Environmental Protection AgencyThe U.S. Environmental Protection Agency is an agency of the federal government of the United States charged with protecting human health and the environment, by writing and enforcing regulations based on laws passed by Congress...
and University of FloridaUniversity of FloridaThe University of Florida is an American public land-grant, sea-grant, and space-grant research university located on a campus in Gainesville, Florida. The university traces its historical origins to 1853, and has operated continuously on its present Gainesville campus since September 1906...
/Institute of Food and Agricultural SciencesInstitute of Food and Agricultural SciencesThe University of Florida’s Institute of Food and Agricultural Sciences is a federal-state-county partnership dedicated to developing knowledge in agriculture, human and natural resources, and the life sciences, and enhancing and sustaining the quality of human life by making that information...
National Public Health Pesticide Applicator Training Manual - Arthropods - Arthropoda Insect Life Forms