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Spiders ( order Araneae) are air-breathing arthropods that have eight legs,
chelicerae The chelicerae () are the mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as " jaws", chelicerae may be shaped as either articulated fangs, or similarl ...
with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of
arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegar ...
s and rank seventh in total species diversity among all orders of organisms. Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every land habitat. , 50,356 spider species in 132 families have been recorded by taxonomists. However, there has been debate among scientists about how families should be classified, with over 20 different classifications proposed since 1900. 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. Unlike
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...
s, spiders do not have antennae. In all except the most primitive group, the
Mesothelae The Mesothelae are a suborder of spiders (order Araneae) that includes a single extant family, Liphistiidae, and a number of extinct families. This suborder is thought to form the sister group to all other living spiders, and to retain ancestral ...
, 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 In anatomy, extension is a movement of a joint that increases the angle between two bones or body surfaces at a joint. Extension usually results in straightening of the bones or body surfaces involved. For example, extension is produced by extendin ...
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 web A spider web, spiderweb, spider's web, or cobweb (from the archaic word '' coppe'', meaning "spider") is a structure created by a spider out of proteinaceous spider silk extruded from its spinnerets, generally meant to catch its prey. Spi ...
s 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 spider Orb-weaver spiders are members of the spider family (biology), family Araneidae. They are the most common group of builders of spiral wheel-shaped spider web, webs often found in gardens, fields, and forests. The English word "orb" can mean "circ ...
s. Spider-like
arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegar ...
s with silk-producing spigots appeared in the Devonian
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
about , but these animals apparently lacked spinnerets. True spiders have been found in Carboniferous rocks from , and are very similar to the most primitive surviving suborder, the
Mesothelae The Mesothelae are a suborder of spiders (order Araneae) that includes a single extant family, Liphistiidae, and a number of extinct families. This suborder is thought to form the sister group to all other living spiders, and to retain ancestral ...
. The main groups of modern spiders, Mygalomorphae and
Araneomorphae The Araneomorphae (also called the Labidognatha) are an infraorder of spiders. They are distinguishable by chelicerae (fangs) that point diagonally forward and cross in a pinching action, in contrast to the Mygalomorphae (tarantulas and their ...
, first appeared in the Triassic period, before . The species ''
Bagheera kiplingi ''Bagheera kiplingi'' is a species of jumping spider found in Central America, including Mexico, Costa Rica, and Guatemala. It is the type species of the genus ''Bagheera'', which includes three other species, including '' B. prosper''.Platnick, ...
'' was described as
herbivorous A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
in 2008, 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. It is estimated that the world's 25 million tons of spiders kill 400–800 million tons of prey per year. Spiders use a wide range of strategies to capture prey: trapping it in sticky webs, lassoing it with sticky
bolas Bolas or bolases (singular bola; from Spanish and Portuguese ''bola'', "ball", also known as a ''boleadora'' or ''boleadeira'') is a type of throwing weapon made of weights on the ends of interconnected cords, used to capture animals by entan ...
, 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, so they liquefy their food by flooding it with digestive
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
s. They also grind food with the bases of their
pedipalp Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") an ...
s, as arachnids do not have the mandibles that crustaceans and insects have. 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 The Mygalomorphae, or mygalomorphs, are an infraorder of spiders, and comprise one of three major groups of living spiders with over 3000 species, found on all continents except Antarctica. Many members are known as trapdoor spiders due to t ...
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.


Etymology

The word ''spider'' derives from Proto-Germanic , literally (a reference to how spiders make their webs), from the Proto-Indo-European root .


Description


Body plan

Spiders are chelicerates and therefore arthropods. 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. 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 Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...
would form two separate tagmata, the head and thorax; the rear tagma is called the abdomen or opisthosoma. In spiders, the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel. 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 The chelicerae () are the mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as " jaws", chelicerae may be shaped as either articulated fangs, or similarl ...
, and they lack anything that would function directly as "jaws". The first appendages behind the mouth are called
pedipalp Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") an ...
s, and serve different functions within different groups of chelicerates. Spiders and scorpions are members of one chelicerate group, the
arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegar ...
s. Scorpions' chelicerae have three sections and are used in feeding. 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. Scorpions' pedipalps generally form large claws for capturing prey, 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. 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 Convex or convexity may refer to: Science and technology * Convex lens, in optics Mathematics * Convex set, containing the whole line segment that joins points ** Convex polygon, a polygon which encloses a convex set of points ** Convex polytop ...
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 The Mesothelae are a suborder of spiders (order Araneae) that includes a single extant family, Liphistiidae, and a number of extinct families. This suborder is thought to form the sister group to all other living spiders, and to retain ancestral ...
, whose living members are the Liphistiidae, have segmented plates on the upper surface.


Circulation and respiration

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. 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 An artery (plural arteries) () is a blood vessel in humans and most animals that takes blood away from the heart to one or more parts of the body (tissues, lungs, brain etc.). Most arteries carry oxygenated blood; the two exceptions are the pu ...
that pass through the pedicle and open into several parts of the cephalothorax. Hence spiders have open circulatory systems. The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as ...
transport more efficient. Spiders have developed several different respiratory anatomies, based on book lungs, a tracheal system, or both.
Mygalomorph The Mygalomorphae, or mygalomorphs, are an infraorder of spiders, and comprise one of three major groups of living spiders with over 3000 species, found on all continents except Antarctica. Many members are known as trapdoor spiders due to t ...
and
Mesothelae The Mesothelae are a suborder of spiders (order Araneae) that includes a single extant family, Liphistiidae, and a number of extinct families. This suborder is thought to form the sister group to all other living spiders, and to retain ancestral ...
spiders have two pairs of book lungs filled with haemolymph, where openings on the
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
surface of the abdomen allow air to enter and diffuse
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as ...
. This is also the case for some basal
araneomorph The Araneomorphae (also called the Labidognatha) are an infraorder of spiders. They are distinguishable by chelicerae (fangs) that point diagonally forward and cross in a pinching action, in contrast to the Mygalomorphae (tarantulas and their ...
spiders, like the family
Hypochilidae Lampshade spiders, family Hypochilidae, are among the most primitive of araneomorph spiders. There are two genera and twelve species currently recognized. Like mygalomorphs, most hypochilids have two pairs of book lungs, but like araneomorphs the ...
, 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. The tracheal system has most likely evolved in small ancestors to help resist desiccation. 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. Spiders that have tracheae generally have higher metabolic rates and better water conservation. Spiders are ectotherms, so environmental temperatures affect their activity.


Feeding, digestion and excretion

Uniquely among chelicerates, the final sections of spiders'
chelicerae The chelicerae () are the mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as " jaws", chelicerae may be shaped as either articulated fangs, or similarl ...
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. The families
Uloboridae Uloboridae is a family of non-venomous spiders, known as cribellate orb weavers or hackled orb weavers. Their lack of venom glands is a secondarily evolved trait. Instead, they wrap their prey thoroughly in silk, cover it in regurgitated digestive ...
and Holarchaeidae, and some Liphistiidae spiders, have lost their venom glands, and kill their prey with silk instead. Like most
arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegar ...
s, including scorpions, spiders have a narrow gut that can only cope with liquid food and two sets of filters to keep solids out. They use one of two different systems of external digestion. Some pump digestive
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
s 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
pedipalp Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") an ...
s, 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. The stomach in the cephalothorax acts as a pump that sends the food deeper into the digestive system. The midgut 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. Most spiders convert
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
ous waste products into
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 a product of the metabolic breakdown ...
, 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. 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, for example the tubules of
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...
s and arachnids develop from completely different parts of the embryo. However, a few primitive spiders, the sub order
Mesothelae The Mesothelae are a suborder of spiders (order Araneae) that includes a single extant family, Liphistiidae, and a number of extinct families. This suborder is thought to form the sister group to all other living spiders, and to retain ancestral ...
and infraorder Mygalomorphae, retain the ancestral arthropod
nephridia The nephridium (plural ''nephridia'') is an invertebrate organ, found in pairs and performing a function similar to the vertebrate kidneys (which originated from the chordate nephridia). Nephridia remove metabolic wastes from an animal's body. Neph ...
("little kidneys"), which use large amounts of water to excrete nitrogenous waste products as ammonia.


Central nervous system

The basic arthropod
central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all p ...
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 The esophagus ( American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to ...
is encircled by this conglomeration of ganglia. Except for the primitive
Mesothelae The Mesothelae are a suborder of spiders (order Araneae) that includes a single extant family, Liphistiidae, and a number of extinct families. This suborder is thought to form the sister group to all other living spiders, and to retain ancestral ...
, 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; in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfused. Despite the relatively small central nervous system, some spiders (like '' Portia'') exhibit complex behaviour, including the ability to use a trial-and-error approach.


Sense organs


Eyes

Spiders have primarily four pairs of eyes on the top-front area of the cephalothorax, arranged in patterns that vary from one family to another. The principal pair at the front are of the type called pigment-cup
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
("little eyes"), which in most arthropods are only capable of detecting the direction from which light is coming, using the shadow cast by the walls of the cup. However, in spiders these eyes are capable of forming images. The other pairs, called secondary eyes, are thought to be derived from the
compound eye A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which disti ...
s of the ancestral chelicerates, but no longer have the separate facets typical of compound eyes. Unlike the principal eyes, in many spiders these secondary eyes detect light reflected from a reflective
tapetum lucidum The ''tapetum lucidum'' ( ; ; ) is a layer of tissue in the eye of many vertebrates and some other animals. Lying immediately behind the retina, it is a retroreflector. It reflects visible light back through the retina, increasing the light a ...
, and wolf spiders can be spotted by torchlight reflected from the tapeta. On the other hand, the secondary eyes of jumping spiders have no tapeta. Other differences between the principal and secondary eyes are that the latter have rhabdomeres that point away from incoming light, just like in vertebrates, while the arrangement is the opposite in the former. The principal eyes are also the only ones with eye muscles, allowing them to move the retina. Having no muscles, the secondary eyes are immobile. The visual acuity of some jumping spiders exceeds by a factor of ten that of
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
, which have by far the best vision among
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...
s. This acuity is achieved by a telephotographic series of lenses, a four-layer retina, and the ability to swivel the eyes and integrate images from different stages in the scan. The downside is that the scanning and integrating processes are relatively slow. There are spiders with a reduced number of eyes, the most common having six eyes (example, '' Periegops suterii'') with a pair of eyes absent on the anterior median line. Other species have four eyes and members of the Caponiidae family can have as few as two. Cave dwelling species have no eyes, or possess vestigial eyes incapable of sight.


Other senses

As with other arthropods, spiders' cuticles 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, spiders and other arthropods have modified their cuticles into elaborate arrays of sensors. Various touch sensors, mostly bristles called setae, respond to different levels of force, from strong contact to very weak air currents. Chemical sensors provide equivalents of taste and smell, often by means of setae. An adult '' Araneus'' may have up to 1,000 such chemosensitive setae, most on the tarsi of the first pair of legs. Males have more chemosensitive bristles on their pedipalps than females. They have been shown to be responsive to sex
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s produced by females, both contact and air-borne. The jumping spider '' Evarcha culicivora'' uses the scent of blood from mammals and other vertebrates, which is obtained by capturing blood-filled
mosquito Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small flies within the family Culicidae (from the Latin ''culex'' meaning " gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish for "li ...
es, to attract the opposite sex. Because they are able to tell the sexes apart, it is assumed the blood scent is mixed with pheromones. Spiders also have in the joints of their limbs slit sensillae that detect force and vibrations. In web-building spiders, all these mechanical and chemical sensors are more important than the eyes, while the eyes are most important to spiders that hunt actively. Like most arthropods, spiders lack balance and acceleration sensors and rely on their eyes to tell them which way is up. Arthropods'
proprioceptor Proprioception ( ), also referred to as kinaesthesia (or kinesthesia), is the sense of self-movement, force, and body position. It is sometimes described as the "sixth sense". Proprioception is mediated by proprioceptors, mechanosensory neurons ...
s, sensors that report the force exerted by muscles and the degree of bending in the body and joints, are well-understood. On the other hand, little is known about what other internal sensors spiders or other arthropods may have. Some spiders use their webs for hearing, where the giant webs function as extended and reconfigurable auditory sensors.


Locomotion

Each of the eight legs of a spider consists of seven distinct parts. The part closest to and attaching the leg to the cephalothorax is the coxa; the next segment is the short
trochanter A trochanter is a tubercle of the femur near its joint with the hip bone. In humans and most mammals, the trochanters serve as important muscle attachment sites. Humans are known to have three trochanters, though the anatomic "normal" includes ...
that works as a hinge for the following long segment, the femur; next is the spider's knee, the patella, which acts as the hinge for the tibia; the
metatarsus The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the me ...
is next, and it connects the tibia to the tarsus (which may be thought of as a foot of sorts); the tarsus ends in a claw made up of either two or three points, depending on the family to which the spider belongs. Although all arthropods use muscles attached to the inside of the exoskeleton to flex their limbs, spiders and a few other groups still use hydraulic pressure to extend them, a system inherited from their pre-arthropod ancestors. The only
extensor In anatomy, extension is a movement of a joint that increases the angle between two bones or body surfaces at a joint. Extension usually results in straightening of the bones or body surfaces involved. For example, extension is produced by extendin ...
muscles in spider legs are located in the three hip joints (bordering the coxa and the trochanter). As a result, a spider with a punctured cephalothorax cannot extend its legs, and the legs of dead spiders curl up. Spiders can generate pressures up to eight times their resting level to extend their legs, and jumping spiders can jump up to 50 times their own length by suddenly increasing the blood pressure in the third or fourth pair of legs. Although larger spiders use hydraulics to straighten their legs, unlike smaller jumping spiders they depend on their
flexor A flexor is a muscle that flexes a joint. In anatomy, flexion (from the Latin verb ''flectere'', to bend) is a joint movement that decreases the angle between the bones that converge at the joint. For example, one’s elbow joint flexes when one ...
muscles to generate the propulsive force for their jumps. Most spiders that hunt actively, rather than relying on webs, have dense tufts of fine bristles between the paired claws at the tips of their legs. These tufts, known as scopulae, consist of bristles whose ends are split into as many as 1,000 branches, and enable spiders with scopulae to walk up vertical glass and upside down on ceilings. It appears that scopulae get their grip from contact with extremely thin layers of water on surfaces. Spiders, like most other
arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegar ...
s, keep at least four legs on the surface while walking or running.


Silk production

The abdomen has no appendages except those that have been modified to form one to four (usually three) pairs of short, movable spinnerets, which emit silk. Each spinneret has many spigots, each of which is connected to one silk gland. There are at least six types of silk gland, each producing a different type of silk. Spitting spiders also produce silk in modified venom glands. Silk is mainly composed of a
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
very similar to that used in insect silk. It is initially a liquid, and hardens not by exposure to air but as a result of being drawn out, which changes the internal structure of the protein. It is similar in tensile strength to nylon and biological materials such as chitin, collagen and
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell w ...
, but is much more elastic. In other words, it can stretch much further before breaking or losing shape. Some spiders have a
cribellum Cribellum literally means "little sieve", and in biology the term generally applies to anatomical structures in the form of tiny perforated plates. In certain groups of diatoms it refers to microscopically punctured regions of the frustule, or o ...
, a modified spinneret with up to 40,000 spigots, each of which produces a single very fine fiber. The fibers are pulled out by the
calamistrum In spiders, the calamistrum is a row of specialized leg bristles used to comb out fine bands of silk. It is only found on cribellate spiders, that is, spiders that possess the spinning organ known as the cribellum. The calamistrum and cribellum ar ...
, a comblike set of bristles on the jointed tip of the cribellum, and combined into a composite woolly thread that is very effective in snagging the bristles of insects. The earliest spiders had cribella, which produced the first silk capable of capturing insects, before spiders developed silk coated with sticky droplets. However, most modern groups of spiders have lost the cribellum. Even species that do not build webs to catch prey use silk in several ways: as wrappers for sperm and for fertilized eggs; as a " safety rope"; for nest-building; and as " parachutes" by the young of some species.


Reproduction and life cycle

Spiders reproduce sexually and fertilization is
internal Internal may refer to: *Internality as a concept in behavioural economics *Neijia, internal styles of Chinese martial arts *Neigong or "internal skills", a type of exercise in meditation associated with Daoism *''Internal (album)'' by Safia, 2016 ...
but indirect, in other words the sperm is not inserted into the female's body by the male's genitals but by an intermediate stage. Unlike many land-living arthropods, male spiders do not produce ready-made spermatophores (packages of sperm), but spin small sperm webs onto which they ejaculate and then transfer the sperm to special syringe-styled structures, palpal bulbs or palpal organs, borne on the tips of the
pedipalp Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") an ...
s of mature males. When a male detects signs of a female nearby he checks whether she is of the same species and whether she is ready to mate; for example in species that produce webs or "safety ropes", the male can identify the species and sex of these objects by "smell". Spiders generally use elaborate courtship rituals to prevent the large females from eating the small males before fertilization, except where the male is so much smaller that he is not worth eating. In web-weaving species, precise patterns of vibrations in the web are a major part of the rituals, while patterns of touches on the female's body are important in many spiders that hunt actively, and may "hypnotize" the female. Gestures and dances by the male are important for
jumping spider Jumping spiders are a group of spiders that constitute the family Salticidae. As of 2019, this family contained over 600 described genera and over 6,000 described species, making it the largest family of spiders at 13% of all species. Jumping spi ...
s, which have excellent eyesight. If courtship is successful, the male injects his sperm from the palpal bulbs into the female via one or two openings on the underside of her abdomen. Female spiders' reproductive tracts are arranged in one of two ways. The ancestral arrangement ("haplogyne" or "non-entelegyne") consists of a single genital opening, leading to two seminal receptacles (spermathecae) in which females store sperm. In the more advanced arrangement ("entelegyne"), there are two further openings leading directly to the spermathecae, creating a "flow through" system rather than a "first-in first-out" one. Eggs are as a general rule only fertilized during oviposition when the stored sperm is released from its chamber, rather than in the ovarian cavity. A few exceptions exist, such as '' Parasteatoda tepidariorum''. In these species the female appears to be able to activate the dormant sperm before oviposition, allowing them to migrate to the ovarian cavity where fertilization occurs. The only known example of direct fertilization between male and female is an Israeli spider, '' Harpactea sadistica'', which has evolved
traumatic insemination Traumatic insemination, also known as hypodermic insemination, is the mating practice in some species of invertebrates in which the male pierces the female's abdomen with his aedeagus and injects his sperm through the wound into her abdominal cav ...
. In this species the male will penetrate its pedipalps through the female's body wall and inject his sperm directly into her ovaries, where the embryos inside the fertilized eggs will start to develop before being laid. Males of the genus ''
Tidarren ''Tidarren'' is a genus of tangle-web spiders first described by Ralph Vary Chamberlin & Wilton Ivie in 1934. Males are much smaller than females, and they amputate one of their palps before maturation, entering their adult life with only one p ...
'' amputate one of their palps before maturation and enter adult life with one palp only. The palps are 20% of the male's body mass in this species, and detaching one of the two improves mobility. In the
Yemen Yemen (; ar, ٱلْيَمَن, al-Yaman), officially the Republic of Yemen,, ) is a country in Western Asia. It is situated on the southern end of the Arabian Peninsula, and borders Saudi Arabia to the Saudi Arabia–Yemen border, north and ...
i species ''Tidarren argo'', the remaining palp is then torn off by the female. The separated palp remains attached to the female's epigynum for about four hours and apparently continues to function independently. In the meantime, the female feeds on the palpless male. In over 60% of cases, the female of the Australian
redback spider The redback spider (''Latrodectus hasselti''), also known as the Australian black widow, is a species of highly venomous spider believed to originate in South Australia or adjacent Western Australian deserts, but now found throughout Australi ...
kills and eats the male after it inserts its second palp into the female's genital opening; in fact, the males co-operate by trying to impale themselves on the females' fangs. Observation shows that most male redbacks never get an opportunity to mate, and the "lucky" ones increase the likely number of offspring by ensuring that the females are well-fed. However, males of most species survive a few matings, limited mainly by their short life spans. Some even live for a while in their mates' webs. DirkvdM Nephila clavipes.jpg, The tiny male of the golden orb weaver ('' Trichonephila clavipes'') (near the top of the leaf) is protected from the female by producing the right vibrations in the web, and may be too small to be worth eating. Orange spider egg sac.jpg, Orange spider egg sac hanging from ceiling Gasteracantha mammosa spiderlings next to their eggs capsule.jpg, '' Gasteracantha mammosa'' spiderlings next to their eggs capsule Lycosidae female carrying young.jpg, Wolf spider carrying its young on its abdomen Females lay up to 3,000 eggs in one or more silk egg sacs, which maintain a fairly constant humidity level. In some species, the females die afterwards, but females of other species protect the sacs by attaching them to their webs, hiding them in nests, carrying them in the
chelicerae The chelicerae () are the mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as " jaws", chelicerae may be shaped as either articulated fangs, or similarl ...
or attaching them to the spinnerets and dragging them along. Baby spiders pass all their
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
l stages inside the egg sac and emerge as spiderlings, very small and sexually immature but similar in shape to adults. Some spiders care for their young, for example a wolf spider's brood clings to rough bristles on the mother's back, and females of some species respond to the "begging" behaviour of their young by giving them their prey, provided it is no longer struggling, or even regurgitate food. Like other arthropods, spiders have to molt to grow as their cuticle ("skin") cannot stretch.Ruppert, 523–24 In some species males mate with newly-molted females, which are too weak to be dangerous to the males. Most spiders live for only one to two years, although some tarantulas can live in captivity for over 20 years, and an Australian female trapdoor spider was documented to have lived in the wild for 43 years, dying of a parasitic wasp attack.


Size

Spiders occur in a large range of sizes. The smallest, '' Patu digua'' from Colombia, are less than in body length. The largest and heaviest spiders occur among tarantulas, which can have body lengths up to and leg spans up to .


Coloration

Only three classes of pigment ( ommochromes, bilins and guanine) have been identified in spiders, although other pigments have been detected but not yet characterized. Melanins, carotenoids and
pterin Pterin is a heterocyclic compound composed of a pteridine ring system, with a "keto group" (a lactam) and an amino group on positions 4 and 2 respectively. It is structurally related to the parent bicyclic heterocycle called pteridine. Pterins, ...
s, very common in other animals, are apparently absent. In some species, the exocuticle of the legs and prosoma is modified by a
tanning Tanning may refer to: *Tanning (leather), treating animal skins to produce leather *Sun tanning, using the sun to darken pale skin **Indoor tanning, the use of artificial light in place of the sun **Sunless tanning, application of a stain or dye t ...
process, resulting in a brown coloration. Bilins are found, for example, in '' Micrommata virescens'', resulting in its green color. Guanine is responsible for the white markings of the European garden spider ''
Araneus diadematus The spider species ''Araneus diadematus'' is commonly called the European garden spider, diadem spider, orangie, cross spider, and crowned orb weaver. It is sometimes called the pumpkin spider, although this name is also used for a different spec ...
''. It is in many species accumulated in specialized cells called guanocytes. In genera such as '' Tetragnatha'', ''
Leucauge ''Leucauge'' () is a spider genus of long-jawed orb weavers, with over 160 species and fully pantropical distribution. The genus was first documented in Scottish zoologist Adam White's 1841 ''Description of new or little known Arachnida''. Cha ...
'', ''
Argyrodes ''Argyrodes'', also called dewdrop spiders, is a genus of comb-footed spiders that was first described by Eugène Louis Simon in 1864. They occur worldwide, and are best known for their kleptoparasitism. They can spin their own webs, but tend to ...
'' or '' Theridiosoma'', guanine creates their silvery appearance. While guanine is originally an end-product of protein metabolism, its excretion can be blocked in spiders, leading to an increase in its storage. Structural colors occur in some species, which are the result of the diffraction, scattering or interference of light, for example by modified setae or scales. The white prosoma of '' Argiope'' results from bristles reflecting the light, ''
Lycosa ''Lycosa'' is a genus of wolf spiders distributed throughout most of the world. Sometimes called the "true tarantula", though not closely related to the spiders most commonly called tarantulas today, ''Lycosa'' spp. can be distinguished from comm ...
'' and ''
Josa The ''Journal of the Optical Society of America'' is a peer-reviewed scientific journal of optics, published by Optica. It was established in 1917 and in 1984 was split into two parts, A and B. ''Journal of the Optical Society of America A'' P ...
'' both have areas of modified cuticle that act as light reflectors. The peacock spiders of Australia (genus '' Maratus'') are notable for their bright structural colours in the males. While in many spiders color is fixed throughout their lifespan, in some groups, color may be variable in response to environmental and internal conditions. Choice of prey may be able to alter the color of spiders. For example, the abdomen of ''
Theridion grallator ''Theridion grallator'', also known as the Hawaiian happy-face spider, is a spider in the family Theridiidae that resides on the Hawaiian Islands. ''T. grallator'' gets its vernacular name of "Hawaiian happy-face spider" from the unique patterns ...
'' will become orange if the spider ingests certain species of Diptera and adult Lepidoptera, but if it consumes
Homoptera Homoptera is a suborder of order Hemiptera that is considered by some taxonomists to be paraphyletic, and therefore deprecated (obsolete). It was therefore split into the suborders Sternorrhyncha, Auchenorrhyncha, and Coleorrhyncha. The earlier w ...
or larval Lepidoptera, then the abdomen becomes green. Environmentally induced color changes may be morphological (occurring over several days) or physiological (occurring near instantly). Morphological changes require pigment synthesis and degradation. In contrast to this, physiological changes occur by changing the position of pigment-containing cells. An example of morphological color changes is background matching. ''
Misumena vatia ''Misumena vatia'' is a species of crab spider with a holarctic distribution. In North America, it is called the goldenrod crab spider or flower (crab) spider,Acorn, John and Sheldon, Ian. (2003). ''Bugs of Ontario'' Edmonton: Lone Pine Publish ...
'' for instance can change its body color to match the substrate it lives on which makes it more difficult to be detected by prey. An example of physiological color change is observed in '' Cyrtophora cicatrosa'', which can change its body color from white to brown near instantly.


Ecology and behavior


Non-predatory feeding

Although spiders are generally regarded as predatory, the
jumping spider Jumping spiders are a group of spiders that constitute the family Salticidae. As of 2019, this family contained over 600 described genera and over 6,000 described species, making it the largest family of spiders at 13% of all species. Jumping spi ...
''
Bagheera kiplingi ''Bagheera kiplingi'' is a species of jumping spider found in Central America, including Mexico, Costa Rica, and Guatemala. It is the type species of the genus ''Bagheera'', which includes three other species, including '' B. prosper''.Platnick, ...
'' gets over 90% of its food from fairly solid plant material produced by acacias as part of a mutually beneficial relationship with a species of
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
. Juveniles of some spiders in the families Anyphaenidae,
Corinnidae Corinnidae is a family of araneomorph spiders, sometimes called corinnid sac spiders. The family, like other "clubionoid" families, has a confusing taxonomic history. Once it was a part of the large catch-all taxon Clubionidae, now very much small ...
, Clubionidae, Thomisidae and
Salticidae Jumping spiders are a group of spiders that constitute the family Salticidae. As of 2019, this family contained over 600 described genera and over 6,000 described species, making it the largest family of spiders at 13% of all species. Jumping spi ...
feed on plant nectar. Laboratory studies show that they do so deliberately and over extended periods, and periodically clean themselves while feeding. These spiders also prefer sugar solutions to plain water, which indicates that they are seeking nutrients. Since many spiders are nocturnal, the extent of nectar consumption by spiders may have been underestimated. Nectar contains amino acids, lipids, vitamins and minerals in addition to sugars, and studies have shown that other spider species live longer when nectar is available. Feeding on nectar avoids the risks of struggles with prey, and the costs of producing venom and digestive enzymes. Various species are known to feed on dead arthropods (scavenging), web silk, and their own shed exoskeletons. Pollen caught in webs may also be eaten, and studies have shown that young spiders have a better chance of survival if they have the opportunity to eat pollen. In captivity, several spider species are also known to feed on bananas,
marmalade Marmalade is a fruit preserve made from the juice and peel of citrus fruits boiled with sugar and water. The well-known version is made from bitter orange. It is also made from lemons, limes, grapefruits, mandarins, sweet oranges, bergamot ...
, milk, egg yolk and sausages.


Capturing prey

The best-known method of prey capture is by means of sticky webs. Varying placement of webs allows different species of spider to trap different insects in the same area, for example flat horizontal webs trap insects that fly up from vegetation underneath while flat vertical webs trap insects in horizontal flight. Web-building spiders have poor vision, but are extremely sensitive to vibrations. Females of the water spider ''
Argyroneta The diving bell spider or water spider (''Argyroneta aquatica'') is the only species of spider known to live almost entirely under water. It is the only member of the genus ''Argyroneta''. When out of the water, the spider ranges in colour from m ...
aquatica'' build underwater "diving bell" webs that they fill with air and use for digesting prey, molting, mating and raising offspring. They live almost entirely within the bells, darting out to catch prey animals that touch the bell or the threads that anchor it. A few spiders use the surfaces of lakes and ponds as "webs", detecting trapped insects by the vibrations that these cause while struggling. Net-casting spiders weave only small webs, but then manipulate them to trap prey. Those of the genus ''
Hyptiotes ''Hyptiotes'' is a genus of spiders in the family (biology), family Uloboridae. Feeding ''Hyptiotes'' creates a triangular web and sits at a vertex until it detects vibrations that signify the collision of its prey. At this moment the spider rel ...
'' and the family Theridiosomatidae stretch their webs and then release them when prey strike them, but do not actively move their webs. Those of the family
Deinopidae Deinopidae, also known as net casting spiders, is a family of cribellate spiders first described by Carl Ludwig Koch in 1850. It consists of stick-like elongated spiders that catch prey by stretching a web across their front legs before propelling ...
weave even smaller webs, hold them outstretched between their first two pairs of legs, and lunge and push the webs as much as twice their own body length to trap prey, and this move may increase the webs' area by a factor of up to ten. Experiments have shown that '' Deinopis spinosus'' has two different techniques for trapping prey: backwards strikes to catch flying insects, whose vibrations it detects; and forward strikes to catch ground-walking prey that it sees. These two techniques have also been observed in other deinopids. Walking insects form most of the prey of most deinopids, but one population of '' Deinopis subrufa'' appears to live mainly on tipulid flies that they catch with the backwards strike. Mature female
bolas spider A bolas spider is a member of the orb-weaver spider (family Araneidae) that, instead of spinning a typical orb web, hunts by using one or more sticky "capture blobs" on the end of a silk line, known as a " bolas". By swinging the bolas at flyin ...
s of the genus '' Mastophora'' build "webs" that consist of only a single "trapeze line", which they patrol. They also construct a
bolas Bolas or bolases (singular bola; from Spanish and Portuguese ''bola'', "ball", also known as a ''boleadora'' or ''boleadeira'') is a type of throwing weapon made of weights on the ends of interconnected cords, used to capture animals by entan ...
made of a single thread, tipped with a large ball of very wet sticky silk. They emit chemicals that resemble the
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s of moths, and then swing the bolas at the moths. Although they miss on about 50% of strikes, they catch about the same weight of insects per night as web-weaving spiders of similar size. The spiders eat the bolas if they have not made a kill in about 30 minutes, rest for a while, and then make new bolas. Juveniles and adult males are much smaller and do not make bolas. Instead they release different pheromones that attract moth flies, and catch them with their front pairs of legs. The primitive Liphistiidae, the "trapdoor spiders" of the family Ctenizidae and many tarantulas are
ambush predator Ambush predators or sit-and-wait predators are carnivorous animals that capture or trap prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey ...
s that lurk in burrows, often closed by trapdoors and often surrounded by networks of silk threads that alert these spiders to the presence of prey. Other ambush predators do without such aids, including many crab spiders, and a few species that prey on bees, which see ultraviolet, can adjust their ultraviolet reflectance to match the flowers in which they are lurking. Wolf spiders,
jumping spider Jumping spiders are a group of spiders that constitute the family Salticidae. As of 2019, this family contained over 600 described genera and over 6,000 described species, making it the largest family of spiders at 13% of all species. Jumping spi ...
s, fishing spiders and some crab spiders capture prey by chasing it, and rely mainly on vision to locate prey. Some jumping spiders of the genus '' Portia'' hunt other spiders in ways that seem intelligent, outflanking their victims or luring them from their webs. Laboratory studies show that ''Portia''s instinctive tactics are only starting points for a
trial-and-error Trial and error is a fundamental method of problem-solving characterized by repeated, varied attempts which are continued until success, or until the practicer stops trying. According to W.H. Thorpe, the term was devised by C. Lloyd Morgan (18 ...
approach from which these spiders learn very quickly how to overcome new prey species. However, they seem to be relatively slow "thinkers", which is not surprising, as their brains are vastly smaller than those of mammalian predators. Ant-mimicking spiders face several challenges: they generally develop slimmer abdomens and false "waists" in the cephalothorax to mimic the three distinct regions (tagmata) of an ant's body; they wave the first pair of legs in front of their heads to mimic antennae, which spiders lack, and to conceal the fact that they have eight legs rather than six; they develop large color patches round one pair of eyes to disguise the fact that they generally have eight simple eyes, while ants have two compound eyes; they cover their bodies with reflective bristles to resemble the shiny bodies of ants. In some spider species, males and females mimic different ant species, as female spiders are usually much larger than males. Ant-mimicking spiders also modify their behavior to resemble that of the target species of ant; for example, many adopt a zig-zag pattern of movement, ant-mimicking jumping spiders avoid jumping, and spiders of the genus '' Synemosyna'' walk on the outer edges of leaves in the same way as ''
Pseudomyrmex ''Pseudomyrmex'' is a genus of stinging, wasp-like ants in the subfamily Pseudomyrmecinae. They are large-eyed, slender ants, found mainly in tropical and subtropical regions of the New World. Distribution and habitat ''Pseudomyrmex'' is predom ...
''. Ant mimicry in many spiders and other arthropods may be for protection from predators that hunt by sight, including birds, lizards and spiders. However, several ant-mimicking spiders prey either on ants or on the ants' " livestock", such as
aphid Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...
s. When at rest, the ant-mimicking crab spider '' Amyciaea'' does not closely resemble '' Oecophylla'', but while hunting it imitates the behavior of a dying ant to attract worker ants. After a kill, some ant-mimicking spiders hold their victims between themselves and large groups of ants to avoid being attacked.


Defense

There is strong evidence that spiders' coloration is camouflage that helps them to evade their major predators, birds and
parasitic wasp Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later causi ...
s, both of which have good color vision. Many spider species are colored so as to merge with their most common backgrounds, and some have disruptive coloration, stripes and blotches that break up their outlines. In a few species, such as the Hawaiian happy-face spider, ''
Theridion grallator ''Theridion grallator'', also known as the Hawaiian happy-face spider, is a spider in the family Theridiidae that resides on the Hawaiian Islands. ''T. grallator'' gets its vernacular name of "Hawaiian happy-face spider" from the unique patterns ...
'', several coloration schemes are present in a ratio that appears to remain constant, and this may make it more difficult for predators to recognize the species. Most spiders are insufficiently dangerous or unpleasant-tasting for warning coloration to offer much benefit. However, a few species with powerful venom, large jaws or irritant bristles have patches of warning colors, and some actively display these colors when threatened. Many of the family
Theraphosidae Tarantulas comprise a group of large and often hairy spiders of the family Theraphosidae. , 1,040 species have been identified, with 156 genera. The term "tarantula" is usually used to describe members of the family Theraphosidae, although m ...
, which includes tarantulas and baboon spiders, have
urticating hair Urticating hairs or urticating bristles are one of the primary defense mechanisms used by numerous plants, almost all New World tarantulas, and various lepidopteran caterpillars. ''Urtica'' is Latin for "nettle" (stinging nettles are in the genu ...
s on their abdomens and use their legs to flick them at attackers. These bristles are fine setae (bristles) with fragile bases and a row of barbs on the tip. The barbs cause intense irritation but there is no evidence that they carry any kind of venom. A few defend themselves against wasps by including networks of very robust threads in their webs, giving the spider time to flee while the wasps are struggling with the obstacles. The golden wheeling spider, '' Carparachne aureoflava'', of the Namibian desert escapes parasitic wasps by flipping onto its side and
cartwheeling A cartwheel is a sideways rotary movement of the body. It is performed by bringing the hands to the floor one at a time while the body inverts. The legs travel over the body trunk while one or both hands are on the floor, and then the feet ret ...
down sand dunes.


Socialization

A few spider species that build webs live together in large colonies and show social behavior, although not as complex as in social insects. '' Anelosimus
eximius Indian Institute of Management Bangalore (IIM Bangalore or IIMB) is a public business school and Institute of National Importance located in Bangalore, India. Founded in 1973, it was the third IIM to be established, after IIM Calcutta and IIM ...
'' (in the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Theridiidae Theridiidae, also known as the tangle-web spiders, cobweb spiders and comb-footed spiders, is a large family of araneomorph spiders first described by Carl Jakob Sundevall in 1833. This diverse, globally distributed family includes over 3,000 sp ...
) can form colonies of up to 50,000 individuals. The genus ''Anelosimus'' has a strong tendency towards sociality: all known American species are social, and species in
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
are at least somewhat social. Members of other species in the same family but several different genera have independently developed social behavior. For example, although ''
Theridion ''Theridion'' is a genus of tangle-web spiders with a worldwide distribution. Notable species are the Hawaiian happy face spider (''T. grallator''), named for the iconic symbol on its abdomen, and '' T. nigroannulatum'', one of few spid ...
nigroannulatum'' belongs to a genus with no other social species, ''T. nigroannulatum'' build colonies that may contain several thousand individuals that co-operate in prey capture and share food. Other communal spiders include several ''
Philoponella ''Philoponella'' is a genus of uloborid spiders. Like all Uloboridae, these species have no venom. Cooperation Some species (among them ''P. congregabilis'' and ''P. oweni'') construct communal webs, but nevertheless do not capture prey cooper ...
'' species (family
Uloboridae Uloboridae is a family of non-venomous spiders, known as cribellate orb weavers or hackled orb weavers. Their lack of venom glands is a secondarily evolved trait. Instead, they wrap their prey thoroughly in silk, cover it in regurgitated digestive ...
), ''
Agelena consociata ''Agelena consociata'' is a social species of funnel web spider that occurs in tropical forests in West Africa and lives in colonies of one to several hundred individuals. This species is found in rainforest habitats in Gabon. It favors dense for ...
'' (family Agelenidae) and '' Mallos gregalis'' (family
Dictynidae Dictynidae is a family of cribellate, hackled band-producing spiders first described by Octavius Pickard-Cambridge in 1871. Most build irregular webs on or near the ground, creating a tangle of silken fibers among several branches or stems of one ...
). Social predatory spiders need to defend their prey against
kleptoparasite Kleptoparasitism (etymologically, parasitism by theft) is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, which can mean when foo ...
s ("thieves"), and larger colonies are more successful in this. The herbivorous spider ''
Bagheera kiplingi ''Bagheera kiplingi'' is a species of jumping spider found in Central America, including Mexico, Costa Rica, and Guatemala. It is the type species of the genus ''Bagheera'', which includes three other species, including '' B. prosper''.Platnick, ...
'' lives in small colonies which help to protect eggs and spiderlings. Even widow spiders (genus ''
Latrodectus ''Latrodectus'' is a broadly distributed genus of spiders with several species that are commonly known as the true widows. This group is composed of those often loosely called black widow spiders, brown widow spiders, and similar spiders. Howeve ...
''), which are notoriously cannibalistic, have formed small colonies in captivity, sharing webs and feeding together. In experiments, spider species like ''
Steatoda grossa ''Steatoda grossa'', commonly known as the cupboard spider, the dark comb-footed spider, the brown house spider (in Australia), or the false widow or false black widow (though several other species are known by these names), is a common species o ...
'', ''
Latrodectus hesperus ''Latrodectus hesperus'', the western black widow spider or western widow, is a venomous spider species found in western regions of North America. The female's body is 14–16 mm (1/2 in) in length and is black, often with an hourglass-sha ...
'' and '' Eratigena agrestis'' stayed away from ''
Myrmica rubra ''Myrmica rubra'', also known as the common red ant or erroneously the European fire ant, is a species of ant of the genus '' Myrmica'', found all over Europe and is now invasive in some parts of North America and Asia. It is mainly red in colou ...
'' ant colonies. These ants are predators and the
pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
they release for communication have a notable deterrent effect on these spider species.


Web types

There is no consistent relationship between the classification of spiders and the types of web they build: species in the same genus may build very similar or significantly different webs. Nor is there much correspondence between spiders' classification and the chemical composition of their silks.
Convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
in web construction, in other words use of similar techniques by remotely related species, is rampant. Orb web designs and the spinning behaviors that produce them are the best understood. The basic radial-then-spiral sequence visible in orb webs and the sense of direction required to build them may have been inherited from the common ancestors of most spider groups. However, the majority of spiders build non-orb webs. It used to be thought that the sticky orb web was an evolutionary innovation resulting in the diversification of the
Orbiculariae Orbiculariae is a potential clade of araneomorph spiders, uniting two groups that make orb webs. Phylogenetic analyses based on morphological characters have generally recovered this clade; analyses based on DNA have regularly concluded that th ...
. Now, however, it appears that non-orb spiders are a subgroup that evolved from orb-web spiders, and non-orb spiders have over 40% more species and are four times as abundant as orb-web spiders. Their greater success may be because sphecid
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. ...
s, which are often the dominant predators of spiders, much prefer to attack spiders that have flat webs.


Orb

About half the potential prey that hit orb webs escape. A web has to perform three functions: intercepting the prey (intersection), absorbing its momentum without breaking (stopping), and trapping the prey by entangling it or sticking to it (retention). No single design is best for all prey. For example: wider spacing of lines will increase the web's area and hence its ability to intercept prey, but reduce its stopping power and retention; closer spacing, larger sticky droplets and thicker lines would improve retention, but would make it easier for potential prey to see and avoid the web, at least during the day. However, there are no consistent differences between orb webs built for use during the day and those built for use at night. In fact, there is no simple relationship between orb web design features and the prey they capture, as each orb-weaving species takes a wide range of prey. The hubs of orb webs, where the spiders lurk, are usually above the center, as the spiders can move downwards faster than upwards. If there is an obvious direction in which the spider can retreat to avoid its own predators, the hub is usually offset towards that direction. Horizontal orb webs are fairly common, despite being less effective at intercepting and retaining prey and more vulnerable to damage by rain and falling debris. Various researchers have suggested that horizontal webs offer compensating advantages, such as reduced vulnerability to wind damage; reduced visibility to prey flying upwards, because of the backlighting from the sky; enabling
oscillation Oscillation is the repetitive or periodic variation, typically in time, of some measure about a central value (often a point of equilibrium) or between two or more different states. Familiar examples of oscillation include a swinging pendulum ...
s to catch insects in slow horizontal flight. However, there is no single explanation for the common use of horizontal orb webs. Spiders often attach highly visible silk bands, called decorations or stabilimenta, to their webs. Field research suggests that webs with more decorative bands captured more prey per hour. However, a laboratory study showed that spiders reduce the building of these decorations if they sense the presence of predators. There are several unusual variants of orb web, many of them convergently evolved, including: attachment of lines to the surface of water, possibly to trap insects in or on the surface; webs with twigs through their centers, possibly to hide the spiders from predators; "ladderlike" webs that appear most effective in catching moths. However, the significance of many variations is unclear. The orb-weaving species, ''
Zygiella x-notata ''Zygiella x-notata'', sometimes known as the missing sector orb weaver or the silver-sided sector spider,Factsheet 6: Missing-sector Orbweaver (Zygiella x-notata)', British Arachnological Society, 2016. Retrieved 27 November 2016 is a spider sp ...
,'' for example, is known for its characteristic missing sector orb web. The missing sector contains a signal thread used to detect prey vibrations on the female's web. In 1973,
Skylab 3 Skylab 3 (also SL-3 and SLM-2) was the second crewed mission to the first American space station, Skylab. The mission began on July 28, 1973, with the launch of NASA astronauts Alan Bean, Owen Garriott, and Jack Lousma in the Apollo command ...
took two orb-web spiders into
space Space is the boundless three-dimensional extent in which objects and events have relative position and direction. In classical physics, physical space is often conceived in three linear dimensions, although modern physicists usually cons ...
to test their web-spinning capabilities in zero gravity. At first, both produced rather sloppy webs, but they adapted quickly.


Cobweb

Members of the family
Theridiidae Theridiidae, also known as the tangle-web spiders, cobweb spiders and comb-footed spiders, is a large family of araneomorph spiders first described by Carl Jakob Sundevall in 1833. This diverse, globally distributed family includes over 3,000 sp ...
weave irregular, tangled, three-dimensional webs, popularly known as cobwebs. There seems to be an evolutionary trend towards a reduction in the amount of sticky silk used, leading to its total absence in some species. The construction of cobwebs is less stereotyped than that of orb-webs, and may take several days.


Other

The
Linyphiidae Linyphiidae, spiders commonly known as sheet weavers (from the shape of their webs), or money spiders (in the United Kingdom, Ireland, Australia, New Zealand, and in Portugal, from the superstition that if such a spider is seen running on you, ...
generally make horizontal but uneven sheets, with tangles of stopping threads above. Insects that hit the stopping threads fall onto the sheet or are shaken onto it by the spider, and are held by sticky threads on the sheet until the spider can attack from below.


Web design in zero gravity

Many experiments have been conducted to study the effect of zero gravity on the design of spider webs. In late 2020, reports of recent experiments were published that indicated that although web design was affected adversely in zero gravity conditions, having access to a light source could orient spiders and enable them to build their normally shaped webs under such conditions.


Evolution


Fossil record

Although the fossil record of spiders is considered poor, almost 1000 species have been described from fossils. Because spiders' bodies are quite soft, the vast majority of fossil spiders have been found preserved in
amber Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In ...
. The oldest known amber that contains fossil arthropods dates from in the Early
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
period. In addition to preserving spiders' anatomy in very fine detail, pieces of amber show spiders mating, killing prey, producing silk and possibly caring for their young. In a few cases, amber has preserved spiders' egg sacs and webs, occasionally with prey attached; the oldest fossil web found so far is 100 million years old. Earlier spider fossils come from a few
lagerstätte A Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These f ...
n, places where conditions were exceptionally suited to preserving fairly soft tissues. The oldest known exclusively terrestrial
arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegar ...
is the
trigonotarbid The order Trigonotarbida is a group of extinct arachnids whose fossil record extends from the late Silurian to the early Permian ( Pridoli to Sakmarian).Dunlop, J. A., Penney, D. & Jekel, D. 2020A summary list of fossil spiders and their relatives ...
'' Palaeotarbus jerami'', from about in the Silurian period, and had a triangular cephalothorax and segmented abdomen, as well as eight legs and a pair of
pedipalp Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") an ...
s. ''
Attercopus ''Attercopus'' is an extinct genus of arachnids, containing one species ''Attercopus fimbriunguis'', known from flattened cuticle fossils from the Panther Mountain Formation in Upstate New York. It is placed in the extinct order Uraraneida, sp ...
fimbriunguis'', from in the Devonian period, bears the earliest known silk-producing spigots, and was therefore hailed as a spider at the time of its discovery. However, these spigots may have been mounted on the underside of the abdomen rather than on spinnerets, which are modified appendages and whose mobility is important in the building of webs. Hence ''Attercopus'' and the similar
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
arachnid ''
Permarachne ''Permarachne'' is an extinct genus of arachnids containing the single species ''Permarachne novokshonovi'' from the Permian (Kungurian) of Russia, found in the Koshelevka Formation near the town of Suksun in Perm Krai. It is closely related to ...
'' may not have been true spiders, and probably used silk for lining nests or producing egg cases rather than for building webs. The largest known fossil spider as of 2011 is the araneid '' Nephila jurassica'', from about , recorded from Daohuogo,
Inner Mongolia Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an autonomous region of the People's Republic of China. Its border includes most of the length of China's border with the country of Mongolia. Inner Mongolia also accounts for a ...
in China. Its body length is almost 25 mm, (i.e., almost one inch). Several Carboniferous spiders were members of the
Mesothelae The Mesothelae are a suborder of spiders (order Araneae) that includes a single extant family, Liphistiidae, and a number of extinct families. This suborder is thought to form the sister group to all other living spiders, and to retain ancestral ...
, a primitive group now represented only by the Liphistiidae. The mesothelid '' Paleothele montceauensis'', from the Late Carboniferous over , had five spinnerets. Although the
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
period saw rapid
diversification Diversification may refer to: Biology and agriculture * Genetic divergence, emergence of subpopulations that have accumulated independent genetic changes * Agricultural diversification involves the re-allocation of some of a farm's resources to ...
of flying
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...
s, there are very few fossil spiders from this period. The main groups of modern spiders, Mygalomorphae and
Araneomorphae The Araneomorphae (also called the Labidognatha) are an infraorder of spiders. They are distinguishable by chelicerae (fangs) that point diagonally forward and cross in a pinching action, in contrast to the Mygalomorphae (tarantulas and their ...
, first appear in the
Triassic The Triassic ( ) is a geologic period and system (stratigraphy), system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Year#Abbreviations yr and ya, Mya), to the beginning of the Jurassic Period 201.36 ...
well before . Some Triassic mygalomorphs appear to be members of the family
Hexathelidae Hexathelidae is a family of mygalomorph spiders. It is one of a number of families and genera of spiders known as funnel-web spiders. In 2018, the family was substantially reduced in size by genera being moved to three separate families: Atracid ...
, whose modern members include the notorious
Sydney funnel-web spider The Sydney funnel-web spider (''Atrax robustus'') is a species of venomous mygalomorph spider native to eastern Australia, usually found within a radius of Sydney. It is a member of a group of spiders known as Australian funnel-web spiders. I ...
, and their spinnerets appear adapted for building funnel-shaped webs to catch jumping insects.
Araneomorphae The Araneomorphae (also called the Labidognatha) are an infraorder of spiders. They are distinguishable by chelicerae (fangs) that point diagonally forward and cross in a pinching action, in contrast to the Mygalomorphae (tarantulas and their ...
account for the great majority of modern spiders, including those that weave the familiar orb-shaped webs. The
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
and
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
periods provide a large number of fossil spiders, including representatives of many modern families.


External relationships

The spiders (Araneae) are monophyletic (i.e., a clade, consisting of a last common ancestor and all of its descendants). There has been debate about what their closest evolutionary relatives are, and how all of these evolved from the ancestral chelicerates, which were marine animals. This 2019 cladogram illustrates the spiders' phylogenetic relationships. Arachnids lack some features of other chelicerates, including backward-pointing mouths and gnathobases ("jaw bases") at the bases of their legs; both of these features are part of the ancestral arthropod feeding system. Instead, they have mouths that point forwards and downwards, and all have some means of breathing air. Spiders (Araneae) are distinguished from other arachnid groups by several characteristics, including spinnerets and, in males,
pedipalp Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") an ...
s that are specially adapted for sperm transfer.


Internal relationships

The
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
shows the relation among spider suborders and families:


Taxonomy

The order name Araneae derives from
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
''aranea'' borrowing
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
''arákhnē'' from ''arákhnēs''. Spiders are divided into two suborders,
Mesothelae The Mesothelae are a suborder of spiders (order Araneae) that includes a single extant family, Liphistiidae, and a number of extinct families. This suborder is thought to form the sister group to all other living spiders, and to retain ancestral ...
and
Opisthothelae The Opisthothelae are spiders within the order Araneae, consisting of the Mygalomorphae and the Araneomorphae, but excluding the Mesothelae. The Opisthothelae are sometimes presented as an unranked clade and sometimes as a suborder of the Arane ...
, of which the latter contains two infraorders, Mygalomorphae and
Araneomorphae The Araneomorphae (also called the Labidognatha) are an infraorder of spiders. They are distinguishable by chelicerae (fangs) that point diagonally forward and cross in a pinching action, in contrast to the Mygalomorphae (tarantulas and their ...
. Some 50,356 living species of spiders (order Araneae) have been identified, grouped into 132
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideal ...
and 4,280 genera by arachnologists in 2022.


Mesothelae

The only living members of the primitive Mesothelae are the family Liphistiidae, found only in
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia, consisting of the regions that are situated south of mainlan ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
, and
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
. Most of the Liphistiidae construct silk-lined burrows with thin
trapdoor A trapdoor is a sliding or hinged door in a floor or ceiling. It is traditionally small in size. It was invented to facilitate the hoisting of grain up through mills, however, its list of uses has grown over time. The trapdoor has played a pivot ...
s, although some species of the genus ''
Liphistius ''Liphistius'' is a genus of basal trapdoor spiders in the family Liphistiidae. They are found in Japan, China, and Southeast Asia. Etymology ''Liphistius'' is from the Greek ''leipo'' (lacking) and ''stios'' (equality). Biology Female body l ...
'' build camouflaged silk tubes with a second trapdoor as an emergency exit. Members of the genus ''Liphistius'' run silk "
tripwire A tripwire is a passive triggering mechanism. Typically, a wire or cord is attached to a device for detecting or reacting to physical movement. Military applications Such tripwires may be attached to one or more mines – especially fragm ...
s" outwards from their tunnels to help them detect approaching prey, while those of the genus ''
Heptathela ''Heptathela'' is a genus of spiders that includes the Kimura spider (''Heptathela kimurai''). They are trapdoor spiders of the family Liphistiidae and are found in Japan, including Okinawa and the Ryukyu Islands. Spiders of this genus lack veno ...
'' do not and instead rely on their built-in vibration sensors. Spiders of the genus ''Heptathela'' have no venom glands, although they do have venom gland outlets on the fang tip. Full text at The extinct families
Arthrolycosidae Arthrolycosidae is an extinct family of arachnids, possibly spiders. Fossils placed in the family were found in the Carboniferous to Permian, . They were considered by Alexander Petrunkevitch to be "mesotheles", i.e. placed in the spider suborde ...
, found in Carboniferous and
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
rocks, and
Arthromygalidae The Arthromygalidae are an extinct family of arachnids, possibly spiders. Fossils placed in the family were all found in the Carboniferous, . They were considered by Alexander Petrunkevitch to be "mesotheles", i.e. placed in the spider suborder M ...
, so far found only in Carboniferous rocks, have been classified as members of the Mesothelae.


Mygalomorphae

The Mygalomorphae, which first appeared in the
Triassic The Triassic ( ) is a geologic period and system (stratigraphy), system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Year#Abbreviations yr and ya, Mya), to the beginning of the Jurassic Period 201.36 ...
period, are generally heavily built and ″hairy″, with large, robust
chelicerae The chelicerae () are the mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as " jaws", chelicerae may be shaped as either articulated fangs, or similarl ...
and fangs (technically, spiders do not have true
hairs Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...
, but rather
setae In biology, setae (singular seta ; from the Latin word for " bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. ...
). Well-known examples include tarantulas, ctenizid trapdoor spiders and the Australasian funnel-web spiders. Most spend the majority of their time in burrows, and some run silk tripwires out from these, but a few build webs to capture prey. However, mygalomorphs cannot produce the pirifom silk that the Araneomorphae use as an instant adhesive to glue silk to surfaces or to other strands of silk, and this makes web construction more difficult for mygalomorphs. Since mygalomorphs rarely "
balloon A balloon is a flexible bag that can be inflated with a gas, such as helium, hydrogen, nitrous oxide, oxygen, and air. For special tasks, balloons can be filled with smoke, liquid water, granular media (e.g. sand, flour or rice), or light so ...
" by using air currents for transport, their populations often form clumps. In addition to arthropods, some mygalomorphs are known to prey on frogs, small mammals, lizards, snakes, snails, and small birds.


Araneomorphae

In addition to accounting for over 90% of spider species, the Araneomorphae, also known as the "true spiders", include orb-web spiders, the
cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often us ...
wolf spiders, and
jumping spider Jumping spiders are a group of spiders that constitute the family Salticidae. As of 2019, this family contained over 600 described genera and over 6,000 described species, making it the largest family of spiders at 13% of all species. Jumping spi ...
s, as well as the only known herbivorous spider, ''
Bagheera kiplingi ''Bagheera kiplingi'' is a species of jumping spider found in Central America, including Mexico, Costa Rica, and Guatemala. It is the type species of the genus ''Bagheera'', which includes three other species, including '' B. prosper''.Platnick, ...
''. They are distinguished by having fangs that oppose each other and cross in a pinching action, in contrast to the Mygalomorphae, which have fangs that are nearly parallel in alignment.


Human interaction


Media coverage and misconceptions

Information about spiders in the media is often emphasizing how dangerous and unpleasant they are. A recent study investigated the global spread of (mis-)information on spiders using a high-resolution global database of online newspaper articles on spider–human interactions. These reports covered stories of spider–human encounters and bites published from 2010–2020. The study found that 47% of articles contained errors and 43% were sensationalist.


Bites

Although spiders are widely feared, only a few species are dangerous to people. Spiders will only bite humans in self-defense, and few produce worse effects than a mosquito bite or bee sting. Most of those with medically serious bites, such as
recluse spider The recluse spiders (''Loxosceles'' (), also known as brown spiders, fiddle-backs, violin spiders, and reapers, is a genus of spiders that was first described by R. T. Lowe in 1832. They are venomous spiders known for their bite, which sometimes ...
s (genus ''Loxosceles'') and widow spiders (genus ''
Latrodectus ''Latrodectus'' is a broadly distributed genus of spiders with several species that are commonly known as the true widows. This group is composed of those often loosely called black widow spiders, brown widow spiders, and similar spiders. Howeve ...
''), would rather flee and bite only when trapped, although this can easily arise by accident. The defensive tactics of Australian funnel-web spiders (family Atracidae) include fang display. Their venom, although they rarely inject much, has resulted in 13 attributed human deaths over 50 years. They have been deemed to be the world's most dangerous spiders on clinical and venom toxicity grounds, though this claim has also been attributed to the Brazilian wandering spider (genus ''
Phoneutria ''Phoneutria'' is a genus of spiders in the family Ctenidae. Phoneutria venom is potentially medically significant to humans. They are mainly found in northern South America, with one species in Central America. Members of the genus are commo ...
''). There were about 100 reliably reported deaths from spider bites in the 20th century, compared to about 1,500 from
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbrell ...
stings. Many alleged cases of spider bites may represent incorrect diagnoses, which would make it more difficult to check the effectiveness of treatments for genuine bites. A review published in 2016 agreed with this conclusion, showing that 78% of 134 published medical case studies of supposed spider bites did not meet the necessary criteria for a spider bite to be verified. In the case of the two genera with the highest reported number of bites, ''Loxosceles'' and ''
Latrodectus ''Latrodectus'' is a broadly distributed genus of spiders with several species that are commonly known as the true widows. This group is composed of those often loosely called black widow spiders, brown widow spiders, and similar spiders. Howeve ...
'', spider bites were not verified in over 90% of the reports. Even when verification had occurred, details of the treatment and its effects were often lacking.


Silk

Because
spider silk Spider silk is a protein fibre spun by spiders. Spiders use their silk to make webs or other structures, which function as sticky nets to catch other animals, or as nests or cocoons to protect their offspring, or to wrap up prey. They can ...
is both light and very strong, attempts are being made to produce it in
goat The goat or domestic goat (''Capra hircus'') is a domesticated species of goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the a ...
s' milk and in the leaves of plants, by means of genetic engineering.


Arachnophobia

Arachnophobia is a
specific phobia Specific phobia is an anxiety disorder, characterized by an extreme, unreasonable, and irrational fear associated with a specific object, situation, or concept which poses little or no actual danger. Specific phobia can lead to avoidance of the o ...
—it is the abnormal fear of spiders or anything reminiscent of spiders, such as webs or spiderlike shapes. It is one of the most common specific phobias, and some statistics show that 50% of women and 10% of men show symptoms. It may be an exaggerated form of an
instinct Instinct is the inherent inclination of a living organism towards a particular complex behaviour, containing both innate (inborn) and learned elements. The simplest example of an instinctive behaviour is a fixed action pattern (FAP), in which a v ...
ive response that helped early humans to survive, or a cultural phenomenon that is most common in predominantly European societies.


As food

Spiders are used as food. Cooked tarantulas are considered a delicacy in
Cambodia Cambodia (; also Kampuchea ; km, កម្ពុជា, UNGEGN: ), officially the Kingdom of Cambodia, is a country located in the southern portion of the Indochinese Peninsula in Southeast Asia, spanning an area of , bordered by Thailan ...
, and by the
Piaroa The Piaroa people, known among themselves as the ''Huottüja'' or ''De'aruhua'', are a pre-Columbian South American indigenous ethnic group of the middle Orinoco Basin in present-day Colombia and Venezuela, living in an area larger than Belgium ...
Indians of southern
Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...
 – provided the highly irritant bristles, the spiders' main defense system, are removed first.


Spiders in culture

Spiders have been the focus of stories and mythologies of various cultures for centuries.
Uttu Uttu was a Mesopotamian goddess of Sumerian religion, Sumerian origin. She was associated with weaving. She appears in multiple myths, such as ''Enki and Ninhursag'' and ''Enki and the World Order''. Name and character Uttu's name was written T ...
, the ancient Sumerian goddess of weaving, was envisioned as a spider spinning her web. According to her main myth, she resisted her father Enki's sexual advances by ensconcing herself in her web, but let him in after he promised her fresh
produce Produce is a generalized term for many farm-produced crops, including fruits and vegetables (grains, oats, etc. are also sometimes considered ''produce''). More specifically, the term ''produce'' often implies that the products are fresh and g ...
as a marriage gift, thereby allowing him to intoxicate her with
beer Beer is one of the oldest and the most widely consumed type of alcoholic drink in the world, and the third most popular drink overall after water and tea. It is produced by the brewing and fermentation of starches, mainly derived from ce ...
and
rape Rape is a type of sexual assault usually involving sexual intercourse or other forms of sexual penetration carried out against a person without their consent. The act may be carried out by physical force, coercion, abuse of authority, or ...
her. Enki's wife
Ninhursag , deity_of=Mother goddess, goddess of fertility, mountains, and rulers , image= Mesopotamian - Cylinder Seal - Walters 42564 - Impression.jpg , caption= Akkadian cylinder seal impression depicting a vegetation goddess, possibly Ninhursag, sittin ...
heard Uttu's screams and rescued her, removing Enki's
semen Semen, also known as seminal fluid, is an organic bodily fluid created to contain spermatozoa. It is secreted by the gonads (sexual glands) and other sexual organs of male or hermaphroditic animals and can fertilize the female ovum. Sem ...
from her
vagina In mammals, the vagina is the elastic, muscular part of the female genital tract. In humans, it extends from the vestibule to the cervix. The outer vaginal opening is normally partly covered by a thin layer of mucosal tissue called the hymen ...
and planting it in the ground to produce eight previously-nonexistent plants. In a story told by the Roman poet
Ovid Pūblius Ovidius Nāsō (; 20 March 43 BC – 17/18 AD), known in English as Ovid ( ), was a Roman poet who lived during the reign of Augustus. He was a contemporary of the older Virgil and Horace, with whom he is often ranked as one of the th ...
in his ''
Metamorphoses The ''Metamorphoses'' ( la, Metamorphōsēs, from grc, μεταμορφώσεις: "Transformations") is a Latin narrative poem from 8 CE by the Roman poet Ovid. It is considered his ''magnum opus''. The poem chronicles the history of the ...
'',
Arachne Arachne (; from , cognate with Latin ) is the protagonist of a tale in Greek mythology known primarily from the version told by the Roman poet Ovid (43 BCE–17 CE), which is the earliest extant source for the story. In Book Six of his ...
(Ancient Greek for "spider") was a Lydian girl who challenged the goddess
Athena Athena or Athene, often given the epithet Pallas, is an ancient Greek religion, ancient Greek goddess associated with wisdom, warfare, and handicraft who was later syncretism, syncretized with the Roman goddess Minerva. Athena was regarded ...
to a weaving contest. Arachne won, but Athena destroyed her tapestry out of jealousy, causing Arachne to hang herself. In an act of mercy, Athena brought Arachne back to life as the first spider. Stories about the
trickster In mythology and the study of folklore and religion, a trickster is a character in a story (god, goddess, spirit, human or anthropomorphisation) who exhibits a great degree of intellect or secret knowledge and uses it to play tricks or otherwi ...
-spider
Anansi Anansi ( ; literally translates to ''spider'') is an Akan folktale character and the Akan God of Stories, Wisdom, Knowledge, and possibly creation. The form of a spider is the most common depiction of Anansi. He is also, sometimes considered t ...
are prominent in the
folktale A folktale or folk tale is a folklore genre that typically consists of a story passed down from generation to generation orally. Folktale may also refer to: Categories of stories * Folkloric tale from oral tradition * Fable (written form of the a ...
s of
West Africa West Africa or Western Africa is the westernmost region of Africa. The United Nations defines Western Africa as the 16 countries of Benin, Burkina Faso, Cape Verde, The Gambia, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia, Mali, M ...
and the Caribbean. In some cultures, spiders have symbolized patience due to their hunting technique of setting webs and waiting for prey, as well as mischief and malice due to their venomous bites. The Italian
tarantella () is a group of various southern Italian folk dances originating in the regions of Calabria, Campania and Puglia. It is characterized by a fast upbeat tempo, usually in time (sometimes or ), accompanied by tambourines. It is among the mo ...
is a dance to rid the young woman of the lustful effects of a spider bite. Web-spinning also caused the association of the spider with creation myths, as they seem to have the ability to produce their own worlds. Dreamcatchers are depictions of spiderwebs. The Moche people of ancient
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = National seal , national_motto = "Firm and Happy f ...
worshipped nature. They placed emphasis on animals and often depicted spiders in their art.Berrin, Katherine & Larco Museum. ''The Spirit of Ancient Peru: Treasures from the Museo Arqueológico Rafael Larco Herrera''. New York:
Thames and Hudson Thames & Hudson (sometimes T&H for brevity) is a publisher of illustrated books in all visually creative categories: art, architecture, design, photography, fashion, film, and the performing arts. It also publishes books on archaeology, history, ...
, 1997.


See also

*
Arachnidism A spider bite, also known as arachnidism, is an injury resulting from the bite of a spider. The effects of most bites are not serious. Most bites result in mild symptoms around the area of the bite. Rarely they may produce a necrotic skin wound ...
* Glossary of spider terms * List of animals that produce silk *
List of endangered spiders An endangered species is "a species at risk of extinction because of human activity, changes in climate, changes in predator-prey ratios, etc., especially when officially designated as such by a governmental agency such as the U.S. Fish and Wildlife ...
*
Spider taxonomy Spider taxonomy is that part of taxonomy that is concerned with the science of naming, defining and classifying all spiders, members of the Araneae order of the arthropod class Arachnida with more than 48,500 described species. However, there a ...
* Toxins


Citations


General and cited references

* *


Further reading

* * * * * * *


External links

*
Picture story about the jumping spider ''Aelurillus v-insignitus''

New Mexico State University "The Spiders of the Arid Southwest"


* [https://web.archive.org/web/20110511234340/http://media.library.uiuc.edu/cgi/b/bib/bix-idx?type=simple&c=bix&sid=54d8a20e4f1eb5f2de074bad4caba7ae&Submit=search&sort=title&q1=spiders&rgn1=Entire+record list of field guides to spiders], from the International Field Guides database *
Record breaking achievements by spiders and the scientists who study them
{{Authority control Articles containing video clips Carboniferous arachnids Extant Pennsylvanian first appearances Taxa named by Carl Alexander Clerck