An arthropod is an invertebrate animal having an exoskeleton (external skeleton), a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda (from Greek ἄρθρον árthron, "joint", and ποδός podós "foot", which together mean "jointed feet"), and include the insects, arachnids, crustaceans, and others. Arthropods are characterized by their jointed limbs and cuticles, 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 appendages. 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 amniotes. 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 brains are formed by fusion of the ganglia of these segments and encircle the esophagus. The respiratory and excretory systems of arthropods vary, depending as much on their environment as on the subphylum to which they belong.
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 spiders are ocelli that can form images and, in a few cases, can swivel to track prey. Arthropods also have a wide range of chemical and mechanical sensors, mostly based on modifications of the many setae (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 scorpions give birth to live young after the eggs have hatched inside the mother. Arthropod hatchlings vary from miniature adults to grubs and caterpillars that lack jointed limbs and eventually undergo a total metamorphosis to produce the adult form. The level of maternal care for hatchlings varies from nonexistent to the prolonged care provided by 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 cycloneuralians (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.