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Spiders (order Araneae) are air-breathing that have eight limbs, chelicerae with generally able to inject venom, and spinnerets that extrude spider silk. They are the largest order of and rank seventh in total species diversity among all orders of . Spiders are found worldwide on every continent except Antarctica, and have become established in nearly every land habitat. , 52,309 spider species in 134 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.
Anatomy, 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 the use of the term "abdomen", as the opisthosoma of all spiders contains a heart and respiratory organs, organs atypical of an abdomen. (2025). 9780764138850, Barron's. ISBN 9780764138850
Unlike , spiders do not have antennae. In all except the most primitive group, the Mesothelae, spiders have the most centralized nervous systems of all arthropods, as all their ganglion are fused into one mass in the cephalothorax. Unlike most arthropods, spiders have no extensor muscles in their limbs and instead extend them by hydraulic pressure.
Their abdomens bear appendages, modified into spinnerets that extrude silk from up to six types of glands. vary widely in size, shape and the amount of sticky thread used. It now appears that the spiral orb web may be one of the earliest forms, and spiders that produce tangled cobwebs are more abundant and diverse than orb-weaver spiders. Spider-like with silk-producing (Uraraneida) appeared in the Devonian period, 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 main groups of modern spiders, Mygalomorphae and Araneomorphae, first appeared in the Triassic period, more than .
The species Bagheera kiplingi was described as Herbivory in 2008, but all other known species are , mostly preying on insects and other spiders, although a few large species also take birds and lizards. An estimated 25 million tons of spiders kill 400–800 million tons of prey every year. Spiders use numerous strategies to capture prey: trapping it in sticky webs, it with sticky bolas, mimicking the prey to avoid detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have acute vision and hunters of the genus Portia show signs of intelligence in their choice of tactics and ability to develop new ones. Spiders' guts are too narrow to take solids, so they liquefy their food by flooding it with digestive . They also grind food with the bases of their , 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 as 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 Latrodectus, to cooperative hunting and food-sharing. Although most spiders live for at most two years, and other Mygalomorphae 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 . Spider silk provides a combination of lightness, strength and elasticity superior to synthetic materials, and spider silk genes have been inserted into and 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.
Spiders and are members of one chelicerate group, the . Scorpions' chelicerae have three sections and are used in feeding. Spiders' chelicerae have two sections and terminate in that are generally , 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, carapace, while the underside is covered by two rather flat plates. The abdomen is soft and egg-shaped. It shows no sign of segmentation, except that the primitive Mesothelae, whose living members are the Liphistiidae, have segmented plates on the upper surface.
Spiders have developed several different respiratory anatomies, based on , a system, or both. Mygalomorph and Mesothelae spiders have two pairs of filled with haemolymph, where openings on the ventral surface of the abdomen allow air to enter and diffuse oxygen. This is also the case for some basal araneomorph spiders, like the family Hypochilidae, but the remaining members of this group have just the anterior pair of book lungs intact while the posterior pair of breathing organs are partly or fully modified into tracheae, through which oxygen is diffused into the haemolymph or directly to the tissue and organs. 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 . Spiders that have tracheae generally have higher and better water conservation. Spiders are , so environmental temperatures affect their activity.
The stomach in the cephalothorax acts as a pump that sends the food deeper into the digestive system. The midgut bears many digestive cecum, 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 waste products into uric acid, which can be excreted as a dry material. Malphigian tubules ("little tubes") extract these wastes from the blood in the hemocoel and dump them into the 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 and arachnids develop from completely different parts of the embryo. However, a few primitive spiders, the suborder Mesothelae and infraorder Mygalomorphae, retain the ancestral arthropod nephridia ("little "), which use large amounts of water to excrete nitrogenous waste products as ammonia.
Despite the relatively small central nervous system, some spiders (like Portia) exhibit complex behaviour, including the ability to use a trial-and-error approach. (1998). 9780120770304, Academic Press. ISBN 9780120770304
Other differences between the principal and secondary eyes are that the latter have Ommatidium 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 dragonfly, which have by far the best vision among . 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 (such as the Kauaʻi cave wolf spider), or possess vestigial eyes incapable of sight (such as Holothele maddeni).
Like most arthropods, spiders lack balance and acceleration sensors and rely on their eyes to tell them which way is up. Arthropods' , 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.Zhou, J., Lai, J., Menda, G., Stafstrom, J.A., Miles, C.I., Hoy, R.R. and Miles, R.N., 2022. Outsourced hearing in an orb-weaving spider that uses its web as an auditory sensor. Proceedings of the National Academy of Sciences, 119(14), p.e2122789119.
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 , keep at least four legs on the surface while walking or running.
Silk is mainly composed of a protein 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, but is much more elastic. In other words, it can stretch much further before breaking or losing shape.
Some spiders have a cribellum, 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, 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 harness"; for nest-building; and as "" by the young of some species.
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 , 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. Complex Genital System of a Haplogyne Spider (Arachnida, Araneae, Tetrablemmidae) Indicates Internal Fertilization and Full Female Control Over Transferred Sperm The only known example of direct fertilization between male and female is an Israeli spider, Harpactea sadistica, which has evolved traumatic insemination. 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 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 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 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.
Females lay up to 3,000 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 or attaching them to the and dragging them along.
Baby spiders pass all their 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. In one exceptional case, females of the jumping spider Toxeus magnus produce a nutritious milk-like substance for their offspring, and fed until they are sexually mature.
Like other , 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 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.
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 will become orange if the spider ingests certain species of Fly and adult Lepidoptera, but if it consumes Homoptera 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 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.
Juveniles of some spiders in the families Anyphaenidae, Corinnidae, Clubionidae, Thomisidae and Salticidae 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 , , 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 , marmalade, milk, egg yolk and . Airborne Spore caught on the webs of orb-weavers may be ingested along with the old web before construction of a new web. The enzyme chitinase present in their digestive fluid allows for the digestion of these spores.
Spiders have been observed to consume plant material belonging to a large variety of taxa and type. Conversely, cursorial spiders comprise the vast majority (over 80%) of reported incidents of plant-eating.
The water spider Argyroneta aquatica build underwater "diving bell" webs that they fill with air and use for digesting prey and molting. Mating and raising the offspring happens in the female's bell. 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 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 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 Tipulidae flies that they catch with the backwards strike.
Mature female of the genus Mastophora build "webs" that consist of only a single "trapeze line", which they patrol. They also construct a bolas made of a single thread, tipped with a large ball of very wet sticky silk. They emit chemicals that resemble the of , 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 Drain fly, and catch them with their front pairs of legs.
The primitive Liphistiidae, the "trapdoor spiders" of the family Ctenizidae and many are 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 , which see ultraviolet, can adjust their ultraviolet reflectance to match the flowers in which they are lurking. , , and some 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 Portias instinctive tactics are only starting points for a trial-and-error 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 mimicry 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. 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 . 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.
Many of the family Theraphosidae, which includes and Harpactirinae, have on their abdomens and use their legs to flick them at attackers. These bristles are fine (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, Wheel spider, of the Namibian desert escapes parasitic wasps by flipping onto its side and cartwheeling down sand .
In experiments, spider species like Steatoda grossa, Latrodectus hesperus and Hobo spider stayed away from Myrmica rubra ant colonies. These ants are predators and the pheromones they release for communication have a notable deterrent effect on these spider species.
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 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 . However, the significance of many variations is unclear. The orb-weaving species, Zygiella x-notata, 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 took two orb-web spiders into space to test their web-spinning capabilities in zero gravity. At first, both produced rather sloppy webs, but they adapted quickly.
The oldest known exclusively terrestrial arachnid is the trigonotarbid 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 . The fossil was originally named Eotarbus but was renamed when it was realized that a Carboniferous arachnid had already been named Eotarbus: Attercopus, 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 , which are modified and whose mobility is important in the building of webs. Hence Attercopus and the similar Permian arachnid Permarachne 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 Araneomorphae Mongolarachne, from about , recorded from Daohuogo, Inner Mongolia in China. Its body length is almost 25 mm, (i.e., almost one inch).
Several Carboniferous spiders were members of the Mesothelae, a primitive group now represented only by the Liphistiidae. The mesothelid Palaeothele montceauensis, from the Late Carboniferous over , had five spinnerets. Although the Permian period saw rapid diversification of flying , there are very few fossil spiders from this period.
The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appear in the Triassic well before . Some Triassic mygalomorphs appear to be members of the family Hexathelidae, whose modern members include the notorious Sydney funnel-web spider, and their spinnerets appear adapted for building funnel-shaped webs to catch jumping insects. Araneomorphae account for the great majority of modern spiders, including those that weave the familiar orb-shaped webs. The Jurassic and Cretaceous periods provide a large number of fossil spiders, including representatives of many modern families.
According to a 2020 study using a molecular clock calibrated with 27 Chelicerata fossils, spiders most likely diverged from other chelicerates between 375 and 328 million years ago.
Arachnids lack some features of other chelicerates, including backward-pointing mouths and ("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 and, in males, that are specially adapted for sperm transfer.
Spiders are divided into two suborders, Mesothelae and Opisthothelae, of which the latter contains two infraorders, Mygalomorphae and Araneomorphae. Some 50,356 living species of spiders (order Araneae) have been identified, grouped into 132 families and 4,280 genus by arachnology in 2022.
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The extinct families Arthrolycosidae, found in Carboniferous and Permian rocks, and Arthromygalidae, so far found only in Carboniferous rocks, have been classified as members of the Mesothelae.
There were about 100 reliably reported deaths from spider bites in the 20th century, compared to about 1,500 from jellyfish 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, 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.
In a story told by the Roman poet Ovid in his Metamorphoses, Arachne (Ancient Greek for "spider") was a Lydian girl who challenged the goddess Athena 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. In a lesser known version of the tale, Athena transformed both Arachne and her brother Phalanx into spiders for committing incest.
Stories about the trickster-spider Anansi are prominent in the oral tradition of West Africa 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 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. are depictions of spiderwebs. The Moche culture people of ancient Peru worshipped nature.Benson, Elizabeth. The Mochica: A Culture of Peru. New York: Praeger Press. 1972. They placed emphasis on animals and often depicted spiders in their art.Berrin, Katherine & Larco Museum. The Spirit of Ancient Peru: Treasures from the Larco Museum. New York: Thames and Hudson, 1997.
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