Spiders (order Araneae) 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 orders of organisms.[1] Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every habitat with the exceptions of air and sea colonization. As of 2008, at least 43,678 spider species,[2] and 109 families have been recorded by taxonomists;[3] however, there has been dissension within the scientific community
as to how all these families should be classified, as evidenced by the
over 20 different classifications that have been proposed since 1900.[4]
Anatomically, spiders differ from other arthropods in that the usual body segments are fused into two tagmata, the cephalothorax and abdomen, and joined by a small, cylindrical pedicel. 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 silk glands within their abdomen. 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-web spiders. Spider-like arachnids with silk-producing spigots 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 order, the Mesothelae. The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appeared in the Triassic period, before 200 million years ago.
A herbivorous species, Bagheera kiplingi, was described in 2008,[5] but all other known species are predators, mostly preying on insects and on other spiders, although a few large species also take birds and lizards. Spiders use a wide range of strategies to capture prey: trapping it in sticky webs, lassoing it with sticky bolas, mimicking the prey to avoid detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have acute vision, and hunters of the genus Portia show signs of intelligence in their choice of tactics and ability to develop new ones. Spiders' guts are too narrow to take solids, and they liquidize their food by flooding it with digestive enzymes and grinding it with the bases of their pedipalps, as they do not have true jaws.
Male spiders identify themselves by a variety of complex courtship rituals to avoid being eaten by the females. 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 abnormal fear of spiders is called arachnophobia.
e basic arthropod central nervous system consists of a pair of nerve cords running below the gut, with paired ganglia as local control centers in all segments; a brain formed by fusion of the ganglia for the head segments ahead of and behind the mouth, so that the esophagus is encircled by this conglomeration of ganglia.[15] Except for the primitive Mesothelae, of which the Liphistiidae are the sole surviving family, spiders have the much more centralized nervous system that is typical of arachnids: all the ganglia of all segments behind the esophagus are fused, so that the cephalothorax is largely filled with nervous tissue and there are no ganglia in the abdomen;[8][9][15] in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfusedSpiders are chelicerates and therefore arthropods.[6] 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.[7] 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.[6] In spiders the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel.[8] 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".[7][9] The first appendages behind the mouth are called pedipalps, and serve different functions within different groups of chelicerates.[6]
Spiders and scorpions are members of one chelicerate group, the arachnids.[9] Scorpions' chelicerae have three sections and are used in feeding.[10] 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.[8] Scorpions' pedipalps generally form large claws for capturing prey,[10] 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.[8]
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.[8]
Like other arthropods, spiders are coelomates in which the coelom is reduced to small areas round 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.[11] 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.[8] The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make oxygen transport more efficient.[9]
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.[8] The trachea system has most likely evolved in small ancestors to help resist desiccation.[9] 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.[8] Spiders that have tracheae generally have higher metabolic rates and better water conservation.[12] Spiders are ectotherms, so environmental temperatures affect their activityUniquely 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.[8] The family Uloboridae has lost its venom glands, and kills its prey with silk instead. Like most arachnids including scorpions,[9] spiders have a narrow gut that can only cope with liquid food and spiders have two sets of filters to keep solids out.[8] 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.[8]
The stomach in the cephalothorax acts as a pump that sends the food deeper into the digestive system. The mid gut bears many digestive ceca, compartments with no other exit, that extract nutrients from the food; most are in the abdomen, which is dominated by the digestive system, but a few are found in the cephalothorax.[8]
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.[8] 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,[14] for example the tubules of insects and arachnids develop from completely different parts of the embryo.[9] However a few primitive spiders, the sub-order Mesothelae and infra-order Mygalomorphae, retain the ancestral arthropod nephridia ("little kidneys"),[8] which use large amounts of water to excrete nitrogenous waste products as ammonia.[14]The basic arthropod central nervous system consists of a pair of nerve cords running below the gut, with paired ganglia as local control centers in all segments; a brain formed by fusion of the ganglia for the head segments ahead of and behind the mouth, so that the esophagus is encircled by this conglomeration of ganglia.[15] Except for the primitive Mesothelae, of which the Liphistiidae are the sole surviving family, spiders have the much more centralized nervous system that is typical of arachnids: all the ganglia of all segments behind the esophagus are fused, so that the cephalothorax is largely filled with nervous tissue and there are no ganglia in the abdomen;[8][9][15] in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfused
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Anatomically, spiders differ from other arthropods in that the usual body segments are fused into two tagmata, the cephalothorax and abdomen, and joined by a small, cylindrical pedicel. 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 silk glands within their abdomen. 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-web spiders. Spider-like arachnids with silk-producing spigots 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 order, the Mesothelae. The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appeared in the Triassic period, before 200 million years ago.
A herbivorous species, Bagheera kiplingi, was described in 2008,[5] but all other known species are predators, mostly preying on insects and on other spiders, although a few large species also take birds and lizards. Spiders use a wide range of strategies to capture prey: trapping it in sticky webs, lassoing it with sticky bolas, mimicking the prey to avoid detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have acute vision, and hunters of the genus Portia show signs of intelligence in their choice of tactics and ability to develop new ones. Spiders' guts are too narrow to take solids, and they liquidize their food by flooding it with digestive enzymes and grinding it with the bases of their pedipalps, as they do not have true jaws.
Male spiders identify themselves by a variety of complex courtship rituals to avoid being eaten by the females. 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 abnormal fear of spiders is called arachnophobia.
e basic arthropod central nervous system consists of a pair of nerve cords running below the gut, with paired ganglia as local control centers in all segments; a brain formed by fusion of the ganglia for the head segments ahead of and behind the mouth, so that the esophagus is encircled by this conglomeration of ganglia.[15] Except for the primitive Mesothelae, of which the Liphistiidae are the sole surviving family, spiders have the much more centralized nervous system that is typical of arachnids: all the ganglia of all segments behind the esophagus are fused, so that the cephalothorax is largely filled with nervous tissue and there are no ganglia in the abdomen;[8][9][15] in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfusedSpiders are chelicerates and therefore arthropods.[6] 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.[7] 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.[6] In spiders the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel.[8] 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".[7][9] The first appendages behind the mouth are called pedipalps, and serve different functions within different groups of chelicerates.[6]
Spiders and scorpions are members of one chelicerate group, the arachnids.[9] Scorpions' chelicerae have three sections and are used in feeding.[10] 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.[8] Scorpions' pedipalps generally form large claws for capturing prey,[10] 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.[8]
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.[8]
Like other arthropods, spiders are coelomates in which the coelom is reduced to small areas round 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.[11] 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.[8] The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make oxygen transport more efficient.[9]
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.[8] The trachea system has most likely evolved in small ancestors to help resist desiccation.[9] 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.[8] Spiders that have tracheae generally have higher metabolic rates and better water conservation.[12] Spiders are ectotherms, so environmental temperatures affect their activityUniquely 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.[8] The family Uloboridae has lost its venom glands, and kills its prey with silk instead. Like most arachnids including scorpions,[9] spiders have a narrow gut that can only cope with liquid food and spiders have two sets of filters to keep solids out.[8] 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.[8]
The stomach in the cephalothorax acts as a pump that sends the food deeper into the digestive system. The mid gut bears many digestive ceca, compartments with no other exit, that extract nutrients from the food; most are in the abdomen, which is dominated by the digestive system, but a few are found in the cephalothorax.[8]
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.[8] 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,[14] for example the tubules of insects and arachnids develop from completely different parts of the embryo.[9] However a few primitive spiders, the sub-order Mesothelae and infra-order Mygalomorphae, retain the ancestral arthropod nephridia ("little kidneys"),[8] which use large amounts of water to excrete nitrogenous waste products as ammonia.[14]The basic arthropod central nervous system consists of a pair of nerve cords running below the gut, with paired ganglia as local control centers in all segments; a brain formed by fusion of the ganglia for the head segments ahead of and behind the mouth, so that the esophagus is encircled by this conglomeration of ganglia.[15] Except for the primitive Mesothelae, of which the Liphistiidae are the sole surviving family, spiders have the much more centralized nervous system that is typical of arachnids: all the ganglia of all segments behind the esophagus are fused, so that the cephalothorax is largely filled with nervous tissue and there are no ganglia in the abdomen;[8][9][15] in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfused
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