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Spider

Spiders (order Araneae) are air-breathing arthropods that have


eight legs, chelicerae with fangs generally able to inject Spiders

venom,[2] and spinnerets that extrude silk.[3] They are the largest Temporal range: Pennsylvanian –
order of arachnids and rank seventh in total species diversity Holocene,
among all orders of organisms.[4][5] Spiders are found worldwide
on every continent except for Antarctica, and have become
established in nearly every land habitat. As of August  2021,
49,623 spider species in 129 families have been recorded by
taxonomists.[1] However, there has been dissension within the
scientific community as to how all these families should be
classified, as evidenced by the over 20 different classifications
that have been proposed since 1900.[6]

Anatomically, spiders (as with all arachnids) differ from other


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

Unlike insects, spiders do not have antennae. In all except the Scientific classification
most primitive group, the Mesothelae, spiders have the most Kingdom: Animalia
centralized nervous systems of all arthropods, as all their ganglia
are fused into one mass in the cephalothorax. Unlike most Phylum: Arthropoda
arthropods, spiders have no extensor muscles in their limbs and Subphylum: Chelicerata
instead extend them by hydraulic pressure.
Class: Arachnida
Their abdomens bear appendages that have been modified into Order: Araneae
spinnerets that extrude silk from up to six types of glands. Spider
Clerck, 1757
webs vary widely in size, shape and the amount of sticky thread
used. It now appears that the spiral orb web may be one of the Suborders
earliest forms, and spiders that produce tangled cobwebs are
more abundant and diverse than orb-weaver spiders. Spider-like Mesothelae

arachnids with silk-producing spigots appeared in the Devonian


period about 386 million years ago, but these animals apparently Opisthothelae

lacked spinnerets. True spiders have been found in Carboniferous


rocks from 318 to 299 million years ago, and are very similar to Mygalomorphae

the most primitive surviving suborder, the Mesothelae. The main Araneomorphae
groups of modern spiders, Mygalomorphae and Araneomorphae,
first appeared in the Triassic period, before 200 million years ago.  See Spider taxonomy.
The species Bagheera kiplingi was described as herbivorous in Diversity[1]
2008,[8] but all other known species are predators, mostly
preying on insects and on other spiders, although a few large 120 families, c. 48,000 species
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.[9] Spiders use a wide range of strategies
to capture prey: trapping it in sticky webs, lassoing it with sticky bolas, mimicking the prey to avoid
detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have
acute vision, and hunters of the genus Portia show signs of intelligence in their choice of tactics and ability to
develop new ones. Spiders' guts are too narrow to take solids, so they liquefy their food by flooding it with
digestive enzymes. They also grind food with the bases of their pedipalps, 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 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.

Contents
Etymology
Description
Body plan
Circulation and respiration
Feeding, digestion and excretion
Central nervous system
Sense organs
Eyes
Other senses
Locomotion
Silk production
Reproduction and life cycle
Size
Coloration
Ecology and behavior
Non-predatory feeding
Capturing prey
Defense
Socialization
Web types
Orb
Cobweb
Other
Web design in zero gravity
Evolution
Fossil record
External relationships
Internal relationships
Taxonomy
Mesothelae
Mygalomorphae
Araneomorphae
Human interaction
Bites
Arachnophobia
As food
Spiders in culture
See also
Footnotes
Bibliography
Further reading
External links

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

Description

Body plan

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

Palystes castaneus female Nos 1 to 14 as for dorsal aspect


dorsal aspect
15: sternum of prosoma
 1: pedipalp 16: pedicel (also called pedicle)
 2: trichobothria 17: book lung sac
  3: carapace of prosoma 18: book lung stigma
(cephalothorax) 19: epigastric fold
 4: opisthosoma (abdomen) 20: epigyne
 5: eyes – AL (anterior lateral) 21: anterior spinneret
    AM (anterior median) 22: posterior spinneret
    PL (posterior lateral)
    PM (posterior median) I, II, III, IV=Leg numbers from anterior to posterior
Leg segments:
 6: coxa
 7: trochanter
 8: femur
 9: patella
10: tibia
11: metatarsus
12: tarsus
13: claw
14: chelicera

chelicerates.[11]

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

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

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.[16] However, in spiders, it occupies only the upper part of the
abdomen, and blood is discharged into the hemocoel by one artery that opens at the rear end of the abdomen
and by branching arteries that pass through the pedicle and open into several parts of the cephalothorax.
Hence spiders have open circulatory systems.[13] The blood of many spiders that have book lungs contains
the respiratory pigment hemocyanin to make oxygen transport more efficient.[14]

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

Feeding, digestion and excretion


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

The stomach in the cephalothorax acts as a pump that sends the food A syrphid fly captured in the web of a
deeper into the digestive system. The midgut bears many digestive spider
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.[13]

Most spiders convert nitrogenous waste products into uric acid,


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

Central nervous system

The basic arthropod central nervous system consists of a pair of nerve cords running below the gut, with
paired ganglia as local control centers in all segments; a brain formed by fusion of the ganglia for the head
segments ahead of and behind the mouth, so that the esophagus is encircled by this conglomeration of
ganglia.[21] Except for the primitive Mesothelae, of which the Liphistiidae are the sole surviving family,
spiders have the much more centralized nervous system that is typical of arachnids: all the ganglia of all
segments behind the esophagus are fused, so that the cephalothorax is largely filled with nervous tissue and
there are no ganglia in the abdomen;[13][14][21] in the Mesothelae, the ganglia of the abdomen and the rear
part of the cephalothorax remain unfused.[17]

Despite the relatively small central nervous system, some spiders (like Portia) exhibit complex behaviour,
including the ability to use a trial-and-error approach.[22][23][24]

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.[13] The principal pair at the front are of the type called
pigment-cup ocelli ("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.[25][26] The other pairs,
called secondary eyes, are thought to be derived from the
compound eyes of the ancestral chelicerates, but no longer have This jumping spider's main ocelli (center
the separate facets typical of compound eyes. Unlike the principal pair) are very acute. The outer pair are
eyes, in many spiders these secondary eyes detect light reflected "secondary eyes" and there are other
from a reflective tapetum lucidum, and wolf spiders can be spotted pairs of secondary eyes on the sides
by torchlight reflected from the tapeta. On the other hand, the and top of its head.[25]
secondary eyes of jumping spiders have no tapeta.[13]

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.[27]

The visual acuity of some jumping spiders exceeds by a factor of


ten that of dragonflies, which have by far the best vision among
insects. This acuity is achieved by a telephotographic series of Eyes of the jumping spider, Plexippus
lenses, a four-layer retina, and the ability to swivel the eyes and paykulli
integrate images from different stages in the scan. The downside is
that the scanning and integrating processes are relatively slow.[22]

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.[28] Other species have four eyes and members
of the Caponiidae family can have as few as two.[29] 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.[25] 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
pheromones produced by females, both contact and air-borne.[30] The jumping spider Evarcha culicivora
uses the scent of blood from mammals and other vertebrates, which is obtained by capturing blood-filled
mosquitoes, to attract the opposite sex. Because they are able to tell the sexes apart, it is assumed the blood
scent is mixed with pheromones.[31] 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.[13]
Like most arthropods, spiders lack balance and acceleration sensors and rely on their eyes to tell them which
way is up. Arthropods' proprioceptors, 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.[25]

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 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 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 Image of a spider leg: 1–coxa; 2–
exoskeleton to flex their limbs, spiders and a few other groups still trochanter; 3–femur; 4–patella; 5–
use hydraulic pressure to extend them, a system inherited from their tibia; 6–metatarsus; 7–tarsus; 8–
pre-arthropod ancestors.[32] The only extensor muscles in spider legs claws
are located in the three hip joints (bordering the coxa and the
trochanter).[33] As a result, a spider with a punctured cephalothorax
cannot extend its legs, and the legs of dead spiders curl up.[13] Spiders can generate pressures up to eight
times their resting level to extend their legs,[34] 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.[13] Although larger spiders use
hydraulics to straighten their legs, unlike smaller jumping spiders they depend on their flexor muscles to
generate the propulsive force for their jumps.[33]

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.[13] Spiders, like most other arachnids, keep at least four legs on the surface while walking or
running.[35]

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.[13]

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.[36] 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.[13]

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.[13]

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.[13]

Reproduction and life cycle

Spiders reproduce sexually and fertilization is internal 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,[37] male spiders do not produce ready-made
spermatophores (packages of sperm), but spin small sperm webs onto An orb weaver producing silk from its
which they ejaculate and then transfer the sperm to special syringe- spinnerets
styled structures, palpal bulbs or palpal organs, borne on the tips of
the pedipalps 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".[13]

Spiders generally use elaborate courtship rituals to prevent the large


females from eating the small males before fertilization, except where
Play media
the male is so much smaller that he is not worth eating. In web-
Mating behaviour of Neriene radiata
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 spiders, 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.[13]

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.[38] 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.[39][40][41] 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.[42]

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 Yemeni species Tidarren argo, the remaining palp is then torn off by the female. The
Spider fertilization systems

Haplogyne or non-entelegyne Entelegyne

Schematic diagrams showing sperm entering and being stored in the spermathecae; eggs leaving the ovaries
and being fertilized; and finally a fertilized egg leaving the female's body

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.[43] 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.[44] 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.[45]


The tiny male Orange spider Gasteracantha Wolf spider carrying its young on
of the golden egg sac hanging mammosa its abdomen
orb weaver from ceiling spiderlings next
(Trichonephila to their eggs
clavipes) (near capsule
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.

Females lay up to 3,000 eggs in one or more silk egg sacs,[13] which maintain a fairly constant humidity
level.[45] 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
spinnerets and dragging them along.[13]

Baby spiders pass all their larval 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,[13] 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.[45]

Like other arthropods, spiders have to molt to grow as their cuticle ("skin") cannot stretch.[46] In some
species males mate with newly-molted females, which are too weak to be dangerous to the males.[45] Most
spiders live for only one to two years, although some tarantulas can live in captivity for over 20 years,[13][47]
and an Australian female trapdoor spider was documented to have lived in the wild for 43 years, dying of a
parasitic wasp attack.[48]

Size

Spiders occur in a large range of sizes. The smallest, Patu digua from Colombia, are less than 0.37  mm
(0.015  in) in body length. The largest and heaviest spiders occur among tarantulas, which can have body
lengths up to 90 mm (3.5 in) and leg spans up to 250 mm (9.8 in).[49]

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 pterins,
very common in other animals, are apparently absent. In some
species, the exocuticle of the legs and prosoma is modified by a
tanning process, resulting in a brown coloration.[50]
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. It is in many species accumulated
in specialized cells called guanocytes. In genera such as Tetragnatha,
Leucauge, Argyrodes 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.[50] 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 and Josa both
have areas of modified cuticle that act as light reflectors.[50] The Goliath birdeater (Theraphosa
peacock spiders of Australia (genus Maratus) are notable for their blondi), the largest spider
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.[50] 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 Diptera and adult Lepidoptera, but if it consumes Homoptera or larval Lepidoptera, then the
abdomen becomes green.[51] 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.[50] 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.[52] An example of physiological color change is observed in Cyrtophora cicatrosa, which
can change its body color from white to brown near instantly.[50]

Ecology and behavior

Non-predatory feeding

Although spiders are generally regarded as predatory, the jumping


spider Bagheera kiplingi 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.[53]

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
A jumping spider seen in Chennai.
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.[54]

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, milk, egg yolk and sausages.[54]

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.[13]
Crab spider with prey
Females of the water spider Argyroneta 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.[55] A few spiders use the surfaces of
lakes and ponds as "webs", detecting trapped insects by the
vibrations that these cause while struggling.[13]

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 The Phonognatha graeffei or leaf-
prey that it sees. These two techniques have also been observed in curling spider's web serves both as a
other deinopids. Walking insects form most of the prey of most trap and as a way of making its
deinopids, but one population of Deinopis subrufa appears to live home in a leaf.
mainly on tipulid flies that they catch with the backwards strike.[56]

Mature female bolas spiders 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 pheromones 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.[57][58] 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.[59]
The primitive Liphistiidae, the "trapdoor spiders" of the family
Ctenizidae and many tarantulas are ambush predators 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.[17]
Other ambush predators do without such aids, including many crab
spiders,[13] 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.[50] Wolf spiders, jumping spiders, fishing spiders
A trapdoor spider in the genus and some crab spiders capture prey by chasing it, and rely mainly on
Cyclocosmia, an ambush predator vision to locate prey.[13]

Some jumping spiders of the


genus Portia hunt other spiders in ways that seem intelligent,[22]
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 approach from which these spiders learn very quickly
how to overcome new prey species.[60] However, they seem to be
relatively slow "thinkers", which is not surprising, as their brains are
vastly smaller than those of mammalian predators.[22]

Ant-mimicking spiders face Portia uses both webs and


several challenges: they cunning, versatile tactics to
generally develop slimmer overcome prey.[60]
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;
An ant-mimicking jumping spider 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 aphids. 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.[61]

Defense

There is strong evidence that spiders' coloration is camouflage that helps them to evade their major predators,
birds and parasitic wasps, 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, 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.[50][62]
Many of the family Theraphosidae, which includes tarantulas and
baboon spiders, have urticating hairs 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.[63] 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.[64] The
golden wheeling spider, Carparachne aureoflava, of the Namibian
desert escapes parasitic wasps by flipping onto its side and
cartwheeling down sand dunes.[65] Threat display by a Sydney funnel-
web spider (Atrax robustus).

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 (in the family Theridiidae) can form colonies of up to
50,000 individuals.[66] The genus Anelosimus has a strong tendency towards sociality: all known American
species are social, and species in Madagascar are at least somewhat social.[67] Members of other species in
the same family but several different genera have independently developed social behavior. For example,
although Theridion 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.[68]
Other communal spiders include several Philoponella species (family Uloboridae), Agelena consociata
(family Agelenidae) and Mallos gregalis (family Dictynidae).[69] Social predatory spiders need to defend
their prey against kleptoparasites ("thieves"), and larger colonies are more successful in this.[70] The
herbivorous spider Bagheera kiplingi lives in small colonies which help to protect eggs and spiderlings.[53]
Even widow spiders (genus Latrodectus), which are notoriously cannibalistic, have formed small colonies in
captivity, sharing webs and feeding together.[71]

In experiments, spider species like Steatoda grossa, Latrodectus hesperus and Eratigena agrestis stayed away
from Myrmica rubra ant colonies. These ants are predators and pheromones they release for communication
have a notable deterrent effect on these spider species.[72]

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 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 The large orb web of Araneus
direction required to build them may have been inherited from the diadematus (European garden
[73] spider).
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. 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 wasps, which are often the dominant
predators of spiders, much prefer to attack spiders that have flat webs.[74]
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.[73]
Nephila clavata, a golden orb weaver
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.[73]

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 oscillations to catch insects in slow
horizontal flight. However, there is no single explanation for the common use of horizontal orb webs.[73]

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.[75] However, a laboratory study
showed that spiders reduce the building of these decorations if they sense the presence of predators.[76]

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.[73] 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.[77]

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.[78]

Cobweb

Members of the family Theridiidae 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.[74]

A funnel web.
Other
The Linyphiidae 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.[79]

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.[80][81]

Evolution

Fossil record

Although the fossil record of spiders is considered poor,[82] almost 1000 species
have been described from fossils.[83] Because spiders' bodies are quite soft, the
vast majority of fossil spiders have been found preserved in amber.[83] The
oldest known amber that contains fossil arthropods dates from
130 million years ago in the Early Cretaceous period. In addition to preserving
spiders' anatomy in very fine detail, pieces of amber show spiders mating, killing
Spider preserved in prey, producing silk and possibly caring for their young. In a few cases, amber
amber has preserved spiders' egg sacs and webs, occasionally with prey attached;[84]
the oldest fossil web found so far is 100  million years old.[85] Earlier spider
fossils come from a few lagerstätten, places where conditions were exceptionally
suited to preserving fairly soft tissues.[84]

The oldest known exclusively terrestrial arachnid is the trigonotarbid Palaeotarbus jerami, from about
420 million years ago in the Silurian period, and had a triangular cephalothorax and segmented abdomen, as
well as eight legs and a pair of pedipalps.[86] Attercopus fimbriunguis, from 386  million years ago in the
Devonian period, bears the earliest known silk-producing spigots, and was therefore hailed as a spider at the
time of its discovery.[87] 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 arachnid Permarachne may not have been true spiders, and
probably used silk for lining nests or producing egg cases rather than for building webs.[3] The largest known
fossil spider as of 2011 is the araneid Nephila jurassica, from about 165  million years ago, recorded from
Daohuogo, Inner Mongolia in China.[88] 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.[87]
The mesothelid Paleothele montceauensis, from the Late Carboniferous over
299 million years ago, had five spinnerets.[89] Although the Permian period 299 to 251 million years ago saw
rapid diversification of flying insects, there are very few fossil spiders from this period.[87]

The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appear in the Triassic well
before 200  million years ago. 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.[87]
External relationships

The spiders (Araneae) are monophyletic (i.e., a clade, consisting of a last common ancestor and all of its
descendants).[90] 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.[90] This 2019 cladogram illustrates
the spiders' phylogenetic relationships.[91][92]

Arachnids lack some features of other chelicerates, including backward-pointing mouths and gnathobases
("jaw bases") at the bases of their legs;[90] both of these features are part of the ancestral arthropod feeding
system.[93] Instead, they have mouths that point forwards and downwards, and all have some means of
breathing air.[90] Spiders (Araneae) are distinguished from other arachnid groups by several characteristics,
including spinnerets and, in males, pedipalps that are specially adapted for sperm transfer.[94]

  Pycnogonida (sea spiders)



Xiphosura (horseshoe crabs)


†Eurypterida (sea scorpions)

Non‑pulmonates
(ticks, harvestmen, etc)

Chelicerata
  pulmonates

Prosomapoda Scorpiones
   


  Tetrapulmonata   Araneae (spiders)
Arachnida    

Pedipalpi (whip
scorpions, etc)


Internal relationships

The cladogram shows the relation among spider suborders and families:[95]

Mesothelae
Liphistiidae


Atypoidea

Mygalomorphae


Avicularioidea


Hypochilidae

Araneae
  Opisthothelae    
Gradungulidae
     
Austrochiloidea

Araneomorphae  
Austrochilidae
   

   
Haplogynae

Araneoclada


Entelegynae

Taxonomy
Spiders are divided into two suborders, Mesothelae and Opisthothelae, of which the latter contains two
infraorders, Mygalomorphae and Araneomorphae. Over 48,000 living species of spiders (order Araneae)
have been identified and as of 2019 grouped into 120 families and about 4,100 genera by arachnologists.[1]

  Spider diversity[1][94][5] Features


(numbers are approximate)

Segmented Striking
Ganglia
plates on direction
Suborder/Infraorder Families Genera Species
top of
in Spinnerets[96] of
abdomen
abdomen[96] fangs[13]
Four pairs, in
some species
one pair
Mesothelae 1 8 116 Yes Yes Downwards
fused, under
middle of and
abdomen forwards

Opisthothelae:
20 350 2,900 One, two or
Mygalomorphae
Only in three pairs
No From sides
Opisthothelae: some fossils under rear of
96 3,700 44,000 abdomen to center,
Araneomorphae
like pincers

Mesothelae
The only living members of the primitive Mesothelae are the family Liphistiidae,
found only in Southeast Asia, China, and Japan.[94] Most of the Liphistiidae
construct silk-lined burrows with thin trapdoors, although some species of the
genus Liphistius build camouflaged silk tubes with a second trapdoor as an
emergency exit. Members of the genus Liphistius run silk "tripwires" outwards
from their tunnels to help them detect approaching prey, while those of the genus
Ryuthela sasakii, a Heptathela do not and instead rely on their built-in vibration sensors.[98] Spiders
member of the of the genus Heptathela have no venom glands, although they do have venom
Liphistiidae[97] gland outlets on the fang tip.[99]

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.[100]

Mygalomorphae

The Mygalomorphae, which first appeared in the Triassic period,[87]


are generally heavily built and ″hairy″, with large, robust chelicerae
and fangs (technically, spiders do not have true hairs, but rather
setae).[101][94] Well-known examples include tarantulas, ctenizid
trapdoor spiders and the Australasian funnel-web spiders.[13] 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 A Mexican red-kneed tarantula
difficult for mygalomorphs. Since mygalomorphs rarely "balloon" by Brachypelma hamorii
using air currents for transport, their populations often form
clumps.[94] In addition to arthropods, some mygalomorphs are
known to prey on frogs, small mammals, lizards, snakes, snails, and small birds.[102][103]

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 wolf
spiders, and jumping spiders,[94] as well as the only known herbivorous spider,
Bagheera kiplingi.[53] 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.[104]

Leucauge venusta, an
orb-web spider

Human interaction
Bites

Although spiders are widely feared, only a few species are dangerous to people.[105] Spiders will only bite
humans in self-defense, and few produce worse effects than a mosquito bite or bee sting.[106] Most of those
with medically serious bites, such as recluse spiders (genus Loxosceles) and widow spiders (genus
Latrodectus), would rather flee and bite only when trapped, although this can easily arise by
accident.[107][108] 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.[109] They have been deemed to be the world's most dangerous spiders on clinical and venom toxicity
grounds,[105] though this claim has also been attributed to the Brazilian wandering spider (genus
Phoneutria).[110]

There were about 100 reliably reported deaths from spider bites in the 20th century,[111] compared to about
1,500 from jellyfish stings.[112] Many alleged cases of spider bites may represent incorrect diagnoses,[113]
which would make it more difficult to check the effectiveness of treatments for genuine bites.[114] 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.[115]

Because spider silk is both light and very strong, attempts are being made to produce it in goats' milk and in
the leaves of plants, by means of genetic engineering.[116][117]

Arachnophobia

Arachnophobia is a specific phobia—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,[118][119] and some statistics
show that 50% of women and 10% of men show symptoms.[120] It may be an exaggerated form of an
instinctive response that helped early humans to survive,[121] or a cultural phenomenon that is most common
in predominantly European societies.[122]

As food

Spiders are used as food.[123] Cooked tarantulas are considered a


delicacy in Cambodia,[124] and by the Piaroa Indians of southern
Venezuela  – provided the highly irritant bristles, the spiders' main
defense system, are removed first.[125]

Spiders in culture
Cooked tarantulas are considered a
Spiders have been the focus of stories and mythologies of various
[126] delicacy in Cambodia.
cultures for centuries. Uttu, the ancient Sumerian goddess of
weaving, was envisioned as a spider spinning her web.[127][128]
According to her main myth, she resisted her father Enki's sexual advances by ensconcing herself in her
web,[128] but let him in after he promised her fresh produce as a marriage gift,[128] thereby allowing him to
intoxicate her with beer and rape her.[128] Enki's wife Ninhursag heard Uttu's screams and rescued her,[128]
removing Enki's semen from her vagina and planting it in the ground to produce eight previously-nonexistent
plants.[128] In a story told by the Roman poet Ovid in his Metamorphoses, Arachne was a Lydian girl who
challenged the goddess Athena to a weaving contest.[129][130]
Arachne won, but Athena destroyed her tapestry out of
jealousy,[130][131] causing Arachne to hang herself.[130][131] In an
act of mercy, Athena brought Arachne back to life as the first
spider.[130][131] Stories about the trickster-spider Anansi are
prominent in the folktales of West Africa and the Caribbean.[132]

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.[133] The Italian This Moche ceramic depicts a
tarantella is a dance to rid the young woman of the lustful effects of a spider, and dates from around 300
spider bite. Web-spinning also caused the association of the spider CE.
with creation myths, as they seem to have the ability to produce their
own worlds.[134] Dreamcatchers are depictions of spiderwebs. The
Moche people of ancient Peru worshipped nature.[135] They placed emphasis on animals and often depicted
spiders in their art.[136]

See also
Glossary of spider terms
List of endangered spiders
Spider taxonomy
Arachnidism
Toxins
List of animals that produce silk

Footnotes
1. "Currently valid spider genera and species" (http://www.wsc.nmbe.ch/statistics/). World Spider
Catalog. Natural History Museum Bern. Retrieved 2019-07-17.
2. Cushing P.E. (2008) Spiders (Arachnida: Araneae). In: Capinera J.L. (eds) Encyclopedia of
Entomology. Springer, p. 3496. doi:10.1007/978-1-4020-6359-6_4320 (https://doi.org/10.100
7%2F978-1-4020-6359-6_4320).
3. Selden, P.A. & Shear, W.A. (December 2008). "Fossil evidence for the origin of spider
spinnerets" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2634869). PNAS. 105 (52):
20781–85. Bibcode:2008PNAS..10520781S (https://ui.adsabs.harvard.edu/abs/2008PNAS..1
0520781S). doi:10.1073/pnas.0809174106 (https://doi.org/10.1073%2Fpnas.0809174106).
PMC 2634869 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2634869). PMID 19104044 (htt
ps://pubmed.ncbi.nlm.nih.gov/19104044).
4. Sebastin, P.A. & Peter, K.V. (eds.) (2009). Spiders of India (https://books.google.com/books?id
=9oVHO-3ZGx4C&printsec=frontcover). Universities Press/Orient Blackswan. ISBN 978-81-
7371-641-6
5. Dimitrov, Dimitar; Hormiga, Gustavo (7 January 2021). "Spider Diversification Through Space
and Time" (https://doi.org/10.1146/annurev-ento-061520-083414). Annual Review of
Entomology. 66 (1): 225–241. doi:10.1146/annurev-ento-061520-083414 (https://doi.org/10.11
46%2Fannurev-ento-061520-083414). ISSN 0066-4170 (https://www.worldcat.org/issn/0066-4
170). Retrieved 10 December 2021.
6. Foelix, Rainer F. (1996). Biology of Spiders (https://archive.org/details/biologyofspiders00foel_
0). New York: Oxford University Press. p. 3 (https://archive.org/details/biologyofspiders00foel_
0/page/3). ISBN 978-0-19-509593-7.
7. Shultz, Stanley; Shultz, Marguerite (2009). The Tarantula Keeper's Guide. Hauppauge, New
York: Barron's. p. 23. ISBN 978-0-7641-3885-0.
8. Meehan, Christopher J.; Olson, Eric J.; Reudink, Matthew W.; Kyser, T. Kurt; Curry, Robert L.
(2009). "Herbivory in a spider through exploitation of an ant–plant mutualism" (https://doi.org/1
0.1016%2Fj.cub.2009.08.049). Current Biology. 19 (19): R892–93.
doi:10.1016/j.cub.2009.08.049 (https://doi.org/10.1016%2Fj.cub.2009.08.049).
PMID 19825348 (https://pubmed.ncbi.nlm.nih.gov/19825348). S2CID 27885893 (https://api.se
manticscholar.org/CorpusID:27885893).
9. Nyffeler, Martin; Birkhofer, Klaus (14 March 2017). "An estimated 400–800 million tons of prey
are annually killed by the global spider community" (https://www.ncbi.nlm.nih.gov/pmc/articles/
PMC5348567). The Science of Nature. 104 (30): 30. Bibcode:2017SciNa.104...30N (https://ui.
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Bibliography
Deeleman-Reinhold, Christa L. (2001). Forest Spiders of South East Asia: With a Revision of
the Sac and Ground Spiders. Brill Publishers. ISBN 978-9004119598.
Ruppert, E.E.; Fox, R.S.; Barnes, R.D. (2004). Invertebrate Zoology (https://archive.org/details/
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Further reading
Bilger, Burkhard (5 March 2007). "Spider Woman" (http://www.newyorker.com/reporting/2007/0
3/05/070305fa_fact_bilger). The New Yorker. pp. 66–73.
Bristowe, W.S. (1976). The World of Spiders. Taplinger Publishing Company. ISBN 978-0-
8008-8598-4. OCLC 256272177 (https://www.worldcat.org/oclc/256272177).
Crompton, John (1950). The Life of the Spider. New York: Mentor. OCLC 1979220 (https://ww
w.worldcat.org/oclc/1979220).
Hillyard, Paul (1994). The Book of the Spider: From Arachnophobia to the Love of Spiders (htt
ps://archive.org/details/bookofspider00pdhi). New York: Random House. ISBN 978-0-679-
40881-9. OCLC 35231232 (https://www.worldcat.org/oclc/35231232).
Kaston, B.J.; Kaston, Elizabeth (1953). How to Know the Spiders; Pictured-Keys for
Determining the More Common Spiders, with Suggestions for Collecting and Studying Them
(https://archive.org/details/howtoknowspiders0000kast) (1st ed.). Dubuque, Iowa: W.C. Brown
Company. OCLC 628203833 (https://www.worldcat.org/oclc/628203833).
Main, Barbara York (1975). Spiders. Sydney: Collins. ISBN 978-0-00-211443-1.
OCLC 123151744 (https://www.worldcat.org/oclc/123151744).
Wise, David A. (1993). Spiders in Ecological Webs. Cambridge studies in ecology.
Cambridge: Cambridge University Press. ISBN 978-0-521-32547-9. OCLC 25833874 (https://
www.worldcat.org/oclc/25833874).

External links
Spiders (https://curlie.org/Science/Biology/Flora_and_Fauna/Animalia/Arthropoda/Arachnida/
Araneae) at Curlie
Picture story about the jumping spider Aelurillus v-insignitus (http://www.naturbildarchiv-guent
er.de/index.php?id=2343&L=1)
New Mexico State University "The Spiders of the Arid Southwest" (http://aces.nmsu.edu/acad
emics/spiders/)
Online Videos of Jumping Spiders (Salticids) and other arachnids (http://www.rkwalton.com/ju
mp.html)
list of field guides to spiders (https://web.archive.org/web/20110511234340/http://media.librar
y.uiuc.edu/cgi/b/bib/bix-idx?type=simple&c=bix&sid=54d8a20e4f1eb5f2de074bad4caba7ae&
Submit=search&sort=title&q1=spiders&rgn1=Entire+record), from the International Field
Guides database
Spider hunts (https://www.youtube.com/watch?v=78mYtPHRqpk) on YouTube
The Spider World Record: https://peerj.com/articles/3972/

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