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Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom,[2] and spinnerets that extrude silk.[3] They are the largest order of arachnids and rank seventh in total species diversity among all orders of organisms.[4][5] Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every land habitat. As of November 2023[update], 51,673 spider species in 136 families have been recorded by taxonomists.[1] However, there has been debate among scientists about how families should be classified, with over 20 different classifications proposed since 1900.[6]

Anatomically, spiders (as with all arachnids) differ from other arthropods in that the usual body segments are fused into two tagmata, the cephalothorax or prosoma, and the opisthosoma, or abdomen, and joined by a small, cylindrical pedicel. However, as there is currently neither paleontological nor embryological evidence that spiders ever had a separate thorax-like division, there exists an argument against the validity of the term cephalothorax, which means fused cephalon (head) and the thorax. Similarly, arguments can be formed against use of the term abdomen, as the opisthosoma of all spiders contains a heart and respiratory organs, organs atypical of an abdomen.[7]

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Unlike insects, spiders do not have antennae. In all except the most primitive group, the Mesothelae, spiders have the most centralized nervous systems of all arthropods, as all their ganglia are fused into one mass in the cephalothorax. Unlike most arthropods, spiders have no extensor muscles in their limbs and instead extend them by hydraulic pressure.

Their abdomens bear appendages that have been modified into spinnerets that extrude silk from up to six types of glands. Spider webs vary widely in size, shape and the amount of sticky thread used. It now appears that the spiral orb web may be one of the earliest forms, and spiders that produce tangled cobwebs are more abundant and diverse than orb-weaver spiders. Spider-like arachnids with silk-producing spigots (Uraraneida) appeared in the Devonian period about 386 million years ago, but these animals apparently lacked spinnerets. True spiders have been found in Carboniferous rocks from 318 to 299 million years ago, and are very similar to the most primitive surviving suborder, the Mesothelae. The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appeared in the Triassic period, before 200 million years ago.

To avoid being eaten by the females, which are typically much larger, male spiders identify themselves to potential mates by a variety of complex courtship rituals. Males of most species survive a few matings, limited mainly by their short life spans. Females weave silk egg-cases, each of which may contain hundreds of eggs. Females of many species care for their young, for example by carrying them around or by sharing food with them. A minority of species are social, building communal webs that may house anywhere from a few to 50,000 individuals. Social behavior ranges from precarious toleration, as in the widow spiders, to co-operative hunting and food-sharing. Although most spiders live for at most two years, tarantulas and other mygalomorph spiders can live up to 25 years in captivity.

While the venom of a few species is dangerous to humans, scientists are now researching the use of spider venom in medicine and as non-polluting pesticides. Spider silk provides a combination of lightness, strength and elasticity that is superior to that of synthetic materials, and spider silk genes have been inserted into mammals and plants to see if these can be used as silk factories. As a result of their wide range of behaviors, spiders have become common symbols in art and mythology symbolizing various combinations of patience, cruelty and creative powers. An irrational fear of spiders is called arachnophobia.

The word spider derives from Proto-Germanic *spin-ron-, literally 'spinner' (a reference to how spiders make their webs), from the Proto-Indo-European root *(s)pen- 'to draw, stretch, spin'.[10]

Spiders are chelicerates and therefore arthropods.[11] As arthropods they have: segmented bodies with jointed limbs, all covered in a cuticle made of chitin and proteins; heads that are composed of several segments that fuse during the development of the embryo.[12] Being chelicerates, their bodies consist of two tagmata, sets of segments that serve similar functions: the foremost one, called the cephalothorax or prosoma, is a complete fusion of the segments that in an insect would form two separate tagmata, the head and thorax; the rear tagma is called the abdomen or opisthosoma.[11] In spiders, the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel.[13] The pattern of segment fusion that forms chelicerates' heads is unique among arthropods, and what would normally be the first head segment disappears at an early stage of development, so that chelicerates lack the antennae typical of most arthropods. In fact, chelicerates' only appendages ahead of the mouth are a pair of chelicerae, and they lack anything that would function directly as "jaws".[12][14] The first appendages behind the mouth are called pedipalps, and serve different functions within different groups of chelicerates.[11]

Spiders and scorpions are members of one chelicerate group, the arachnids.[14] Scorpions' chelicerae have three sections and are used in feeding.[15] Spiders' chelicerae have two sections and terminate in fangs that are generally venomous, and fold away behind the upper sections while not in use. The upper sections generally have thick "beards" that filter solid lumps out of their food, as spiders can take only liquid food.[13] Scorpions' pedipalps generally form large claws for capturing prey,[15] while those of spiders are fairly small appendages whose bases also act as an extension of the mouth; in addition, those of male spiders have enlarged last sections used for sperm transfer.[13]

In spiders, the cephalothorax and abdomen are joined by a small, cylindrical pedicel, which enables the abdomen to move independently when producing silk. The upper surface of the cephalothorax is covered by a single, convex carapace, while the underside is covered by two rather flat plates. The abdomen is soft and egg-shaped. It shows no sign of segmentation, except that the primitive Mesothelae, whose living members are the Liphistiidae, have segmented plates on the upper surface.[13]

Like other arthropods, spiders are coelomates in which the coelom 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. The heart is a tube in the upper part of the body, with a few ostia that act as non-return valves allowing blood to enter the heart from the hemocoel but prevent it from leaving before it reaches the front end.[16] However, in spiders, it occupies only the upper part of the abdomen, and blood is discharged into the hemocoel by one artery that opens at the rear end of the abdomen and by branching arteries that pass through the pedicle and open into several parts of the cephalothorax. Hence spiders have open circulatory systems.[13] The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make oxygen transport more efficient.[14]

Spiders have developed several different respiratory anatomies, based on book lungs, a tracheal system, or both. Mygalomorph and Mesothelae spiders have two pairs of book lungs filled with haemolymph, where openings on the ventral surface of the abdomen allow air to enter and diffuse oxygen. This is also the case for some basal araneomorph spiders, like the family Hypochilidae, but the remaining members of this group have just the anterior pair of book lungs intact while the posterior pair of breathing organs are partly or fully modified into tracheae, through which oxygen is diffused into the haemolymph or directly to the tissue and organs.[13] The tracheal system has most likely evolved in small ancestors to help resist desiccation.[14] The trachea were originally connected to the surroundings through a pair of openings called spiracles, but in the majority of spiders this pair of spiracles has fused into a single one in the middle, and moved backwards close to the spinnerets.[13] Spiders that have tracheae generally have higher metabolic rates and better water conservation.[17] Spiders are ectotherms, so environmental temperatures affect their activity.[18]

Uniquely among chelicerates, the final sections of spiders' chelicerae are fangs, and the great majority of spiders can use them to inject venom into prey from venom glands in the roots of the chelicerae.[13] The families Uloboridae and Holarchaeidae, and some Liphistiidae spiders, have lost their venom glands, and kill their prey with silk instead.[19] Like most arachnids, including scorpions,[14] spiders have a narrow gut that can only cope with liquid food and two sets of filters to keep solids out.[13] They use one of two different systems of external digestion. Some pump digestive enzymes from the midgut into the prey and then suck the liquified tissues of the prey into the gut, eventually leaving behind the empty husk of the prey. Others grind the prey to pulp using the chelicerae and the bases of the pedipalps, while flooding it with enzymes; in these species, the chelicerae and the bases of the pedipalps form a preoral cavity that holds the food they are processing.[13]

Most spiders convert nitrogenous waste products into uric acid, which can be excreted as a dry material. Malphigian tubules ("little tubes") extract these wastes from the blood in the hemocoel and dump them into the cloacal chamber, from which they are expelled through the anus.[13] Production of uric acid and its removal via Malphigian tubules are a water-conserving feature that has evolved independently in several arthropod lineages that can live far away from water,[20] for example the tubules of insects and arachnids develop from completely different parts of the embryo.[14] However, a few primitive spiders, the suborder Mesothelae and infraorder Mygalomorphae, retain the ancestral arthropod nephridia ("little kidneys"),[13] which use large amounts of water to excrete nitrogenous waste products as ammonia.[20] be457b7860