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The spider, (family Agelenidae) hides inside the mouth of the funnel and waits for unsuspecting prey.

 

Pima County, Arizona.

Agelena labyrinthica is a species of spiders in the family Agelenidae...A. labyrinthica build flat plate surface webs connected to funnel-shaped retreats similar to labyrinths, which are typically constructed between low lying grass and vegetation.[1] These webs can be at ground level, or up to 1.5 metres (4 ft 11 in) from the ground, however, the majority are found approximately 60 centimetres (24 in) off of the ground.These spiders are fairly common in Europe and Central Europe, and are typically concentrated in areas near forests and low lying vegetation, as well as in dry grasslands

My first focus stacking on a live animal.

 

Focus Stacking with 4 shoots.

 

Family: Agelenidae

Genus: Agelena

 

Wikipedia: en.wikipedia.org/wiki/Agelenidae

Gewone Huisspin, Giant housespider, Tegenaria atrica

 

De gewone huisspin is een spinnensoort uit de familie trechterspinnen (Agelenidae). Het mannetje wordt meestal tussen de 6 tot 9 millimeter lang en het vrouwtje tussen de 7 en 11 millimeter. Hun poten zijn 50-60mm long. Hoewel de spin niet zo heel groot is, lijkt hij wel groot door zijn lange poten. De huisspin leeft solitair en is hoofdzakelijk 's nachts actief. Over het algemeen vangt hij een prooidier die op het web terecht komt. Mocht dit niet gebeuren en is hij erg hongerig, zal de huisspin het web verlaten om zelf op jacht te gaan naar een prooidier. Als de spin een prooi eenmaal heeft gevangen wordt deze met een enzymenvloeistof geïnjecteerd en vervolgens leeggezogen.De spin kan wel een leeftijd van zes jaar bereiken. Je kunt deze spinnen het hele jaar door vinden in huizen, schuren, tuinen of onder houtstapels.

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The Giant house spider is a member of the family Agelenidae.

The two sexes do not differ in coloration or markings. Its coloration is mainly dark brown. The male body and head 6-9mm and female 7-11mm. Legs are 50 to 60mm long.

They make a thick sheet web, about 15cm across, usually in a neglected corner of a house or shed. The giant house spider lives single and mainly is active atnight. The web has a tubular retreat at the rear where the spider sits and waits for dinner to drop by. Food: Small insects which are caught in a sheet web. House spiders can survive for several months without food or water. You can find this spider all year round in houses (especially attic spaces), sheds, garages and other out buildings.

 

Gate Keeper: North American Hobo Spider (Agelenidae) with large, flat funnel web, complete with a front entrance and back door exit. Seen in a field on a summer morning, Rocky Mountain Front Range, Colorado.

 

Phylum Arthropoda

Class Arachnida

Family Agelenidae

Genus Eratigena

Species E. agrestis

North American Hobo Spider (Agelenidae) with large, flat funnel web, complete with a front entrance and back door exit. Seen in a field on a summer morning, Rocky Mountain Front Range, Colorado.

 

Phylum Arthropoda

Class Arachnida

Family Agelenidae

Genus Eratigena

Species E. agrestis

Gate Keeper: North American Hobo Spider (Agelenidae) with large, flat funnel web, complete with a front entrance and back door exit. Seen in a field on a summer morning, Rocky Mountain Front Range, Colorado.

 

Phylum Arthropoda

Class Arachnida

Family Agelenidae

Genus Eratigena

Species E. agrestis

Kingdom: Animalia

Phylum : Arthropoda

Class : Arachnida

Order : Araneae

Family : Agelenidae

Genus : Agelena

Species : A. labyrinthica

Spiders (order Araneae) are air-breathing arthropods that have eight legs and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other orders of organisms. Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every habitat with the exceptions of air and sea colonization. As of November 2015, at least 45,700 spider species, and 114 families have been recorded by taxonomists. 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.

 

Anatomically, spiders differ from other arthropods in that the usual body segments are fused into two tagmata, the cephalothorax and abdomen, and joined by a small, cylindrical pedicel. Unlike insects, spiders do not have antennae. In all except the most primitive group, the Mesothelae, spiders have the most centralized nervous systems of all arthropods, as all their ganglia are fused into one mass in the cephalothorax. Unlike most arthropods, spiders have no extensor muscles in their limbs and instead extend them by hydraulic pressure.

 

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

 

A herbivorous species, Bagheera kiplingi, was described in 2008, but all other known species are predators, mostly preying on insects and on other spiders, although a few large species also take birds and lizards. Spiders use a wide range of strategies to capture prey: trapping it in sticky webs, lassoing it with sticky bolas, mimicking the prey to avoid detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have acute vision, and hunters of the genus Portia show signs of intelligence in their choice of tactics and ability to develop new ones. Spiders' guts are too narrow to take solids, and they liquefy their food by flooding it with digestive enzymes and grinding it with the bases of their pedipalps, as they do not have true jaws.

 

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

 

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

 

DESCRIPTION

BODY PLAN

Spiders are chelicerates and therefore arthropods. As arthropods they have: segmented bodies with jointed limbs, all covered in a cuticle made of chitin and proteins; heads that are composed of several segments that fuse during the development of the embryo.[7] Being chelicerates, their bodies consist of two tagmata, sets of segments that serve similar functions: the foremost one, called the cephalothorax or prosoma, is a complete fusion of the segments that in an insect would form two separate tagmata, the head and thorax; the rear tagma is called the abdomen or opisthosoma. In spiders, the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel. The pattern of segment fusion that forms chelicerates' heads is unique among arthropods, and what would normally be the first head segment disappears at an early stage of development, so that chelicerates lack the antennae typical of most arthropods. In fact, chelicerates' only appendages ahead of the mouth are a pair of chelicerae, and they lack anything that would function directly as "jaws". The first appendages behind the mouth are called pedipalps, and serve different functions within different groups of chelicerates.

 

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

 

In spiders, the cephalothorax and abdomen are joined by a small, cylindrical pedicel, which enables the abdomen to move independently when producing silk. The upper surface of the cephalothorax is covered by a single, convex 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.

 

Like other arthropods, spiders are coelomates in which the coelom is reduced to small areas round the reproductive and excretory systems. Its place is largely taken by a hemocoel, a cavity that runs most of the length of the body and through which blood flows. The heart is a tube in the upper part of the body, with a few ostia that act as non-return valves allowing blood to enter the heart from the hemocoel but prevent it from leaving before it reaches the front end. 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. The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make oxygen transport more efficient.

 

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. The trachea system has most likely evolved in small ancestors to help resist desiccation. The trachea were originally connected to the surroundings through a pair of openings called spiracles, but in the majority of spiders this pair of spiracles has fused into a single one in the middle, and moved backwards close to the spinnerets. Spiders that have tracheae generally have higher metabolic rates and better water conservation. Spiders are ectotherms, so environmental temperatures affect their activity.

 

FEEDING, DIGESTION AND EXCRETION

Uniquely among chelicerates, the final sections of spiders' chelicerae are fangs, and the great majority of spiders can use them to inject venom into prey from venom glands in the roots of the chelicerae. The family Uloboridae has lost its venom glands, and kills its prey with silk instead. Like most arachnids, including scorpions, spiders have a narrow gut that can only cope with liquid food and spiders have two sets of filters to keep solids out. 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.

 

The stomach in the cephalothorax acts as a pump that sends the food deeper into the digestive system. The mid gut bears many digestive ceca, compartments with no other exit, that extract nutrients from the food; most are in the abdomen, which is dominated by the digestive system, but a few are found in the cephalothorax.

 

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. Production of uric acid and its removal via Malphigian tubules are a water-conserving feature that has evolved independently in several arthropod lineages that can live far away from water,[14] for example the tubules of insects and arachnids develop from completely different parts of the embryo. However, a few primitive spiders, the sub-order Mesothelae and infra-order Mygalomorphae, retain the ancestral arthropod nephridia ("little kidneys"), which use large amounts of water to excrete nitrogenous waste products as ammonia.

 

CENTRAL NERVOS 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. 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; in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfused.

 

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

 

SENSE ORGANS

EYES

Most spiders have four pairs of eyes on the top-front area of the cephalothorax, arranged in patterns that vary from one family to another. The 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, the main eyes at the front of spiders' heads are pigment-cup ocelli that are capable of forming images. The other eyes are thought to be derived from the compound eyes of the ancestral chelicerates, but no longer have the separate facets typical of compound eyes. Unlike the main eyes, in many spiders these secondary eyes detect light reflected from a reflective tapetum lucidum, and wolf spiders can be spotted by torch light reflected from the tapeta. On the other hand, jumping spiders' secondary eyes have no tapeta. Some jumping spiders' visual acuity exceeds by a factor of ten that of dragonflies, which have by far the best vision among insects; in fact the human eye is only about five times sharper than a jumping spider's. They achieve this by a telephoto-like series of lenses, a four-layer retina and the ability to swivel their 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, of these those with six-eyes are the most numerous and are missing a pair of eyes on the anterior median line, others species have four-eyes and some just two. Cave dwelling species have no eyes, or possess vestigial eyes incapable of sight.

 

OTHER SENSES

As with other arthropods, spiders' cuticles would block out information about the outside world, except that they are penetrated by many sensors or connections from sensors to the nervous system. In fact, spiders and other arthropods have modified their cuticles into elaborate arrays of sensors. Various touch sensors, mostly bristles called setae, respond to different levels of force, from strong contact to very weak air currents. Chemical sensors provide equivalents of taste and smell, often by means of setae. Pedipalps carry a large number of such setae sensitive to contact chemicals and air-borne smells, such as female pheromones. Spiders also have in the joints of their limbs slit sensillae that detect forces and vibrations. In web-building spiders, all these mechanical and chemical sensors are more important than the eyes, while the eyes are most important to spiders that hunt actively.

 

Like most arthropods, spiders lack balance and acceleration sensors and rely on their eyes to tell them which way is up. Arthropods' 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.

 

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 exoskeleton to flex their limbs, spiders and a few other groups still use hydraulic pressure to extend them, a system inherited from their pre-arthropod ancestors. The only extensor muscles in spider legs are located in the three hip joints (bordering the coxa and the trochanter). As a result, a spider with a punctured cephalothorax cannot extend its legs, and the legs of dead spiders curl up. Spiders can generate pressures up to eight times their resting level to extend their legs, and jumping spiders can jump up to 50 times their own length by suddenly increasing the blood pressure in the third or fourth pair of legs. Although larger spiders use hydraulics to straighten their legs, unlike smaller jumping spiders they depend on their flexor muscles to generate the propulsive force for their jumps.

 

Most spiders that hunt actively, rather than relying on webs, have dense tufts of fine hairs 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 arachnids, keep at least four legs on the surface while walking or running.

 

SILK PRODUCTION

The abdomen has no appendages except those that have been modified to form one to four (usually three) pairs of short, movable spinnerets, which emit silk. Each spinneret has many spigots, each of which is connected to one silk gland. There are at least six types of silk gland, each producing a different type of silk.

 

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 comb-like 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.

 

Tarantulas also have silk glands in their feet.

 

Even species that do not build webs to catch prey use silk in several ways: as wrappers for sperm and for fertilized eggs; as a "safety rope"; for nest-building; and as "parachutes" by the young of some species.

 

REPRODUCTION AND LIFE CYCLE

Spiders reproduce sexually and fertilization is internal but indirect, in other words the sperm is not inserted into the female's body by the male's genitals but by an intermediate stage. Unlike many land-living arthropods, male spiders do not produce ready-made spermatophores (packages of sperm), but spin small sperm webs on to which they ejaculate and then transfer the sperm to special syringe-like 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".

 

Spiders generally use elaborate courtship rituals to prevent the large females from eating the small males before fertilization, except where the male is so much smaller that he is not worth eating. In web-weaving species, precise patterns of vibrations in the web are a major part of the rituals, while patterns of touches on the female's body are important in many spiders that hunt actively, and may "hypnotize" the female. Gestures and dances by the male are important for jumping spiders, which have excellent eyesight. If courtship is successful, the male injects his sperm from the palpal bulbs into the female's genital opening, known as the epigyne, on the underside of her abdomen. Female's reproductive tracts vary from simple tubes to systems that include seminal receptacles in which females store sperm and release it when they are ready.

 

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 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 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 eggs in one or more silk egg sacs, which maintain a fairly constant humidity level. In some species, the females die afterwards, but females of other species protect the sacs by attaching them to their webs, hiding them in nests, carrying them in the chelicerae or attaching them to the spinnerets and dragging them along.

 

Baby spiders pass all their larval stages inside the egg and hatch 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 cling to rough bristles on the mother's back, and females of some species respond to the "begging" behaviour of their young by giving them their prey, provided it is no longer struggling, or even regurgitate food.

 

Like other arthropods, spiders have to molt to grow as their cuticle ("skin") cannot stretch. In some species males mate with newly molted females, which are too weak to be dangerous to the males. Most spiders live for only one to two years, although some tarantulas can live in captivity for over 20 years.

 

SIZE

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

 

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.

 

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 amino acids, lipids, vitamins and minerals in addition to sugars, and studies have shown that other spider species live longer when nectar is available. Feeding on nectar avoids the risks of struggles with prey, and the costs of producing venom and digestive enzymes.

 

Various species are known to feed on dead arthropods (scavenging), web silk, and their own shed exoskeletons. Pollen caught in webs may also be eaten, and studies have shown that young spiders have a better chance of survival if they have the opportunity to eat pollen. In captivity, several spider species are also known to feed on bananas, marmalade, milk, egg yolk and sausages.

 

METHODS OF CAPTURING PREY

The best-known method of prey capture is by means of sticky webs. Varying placement of webs allows different species of spider to trap different insects in the same area, for example flat horizontal webs trap insects that fly up from vegetation underneath while flat vertical webs trap insects in horizontal flight. Web-building spiders have poor vision, but are extremely sensitive to vibrations.

 

Females of the water spider Argyroneta aquatica build underwater "diving bell" webs that they fill with air and use for digesting prey, molting, mating and raising offspring. They live almost entirely within the bells, darting out to catch prey animals that touch the bell or the threads that anchor it. A few spiders use the surfaces of lakes and ponds as "webs", detecting trapped insects by the vibrations that these cause while struggling.

 

Net-casting spiders weave only small webs, but then manipulate them to trap prey. Those of the genus Hyptiotes 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 tipulid flies that they catch with the backwards strike.

 

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. Juveniles and adult males are much smaller and do not make bolas. Instead they release different pheromones that attract moth flies, and catch them with their front pairs of legs.

 

The primitive Liphistiidae, the "trapdoor spiders" of the family Ctenizidae and many tarantulas are ambush 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. Other ambush predators do without such aids, including many crab spiders, and a few species that prey on bees, which see ultraviolet, can adjust their ultraviolet reflectance to match the flowers in which they are lurking. Wolf spiders, jumping spiders, fishing spiders and some crab spiders capture prey by chasing it, and rely mainly on vision to locate prey.Some jumping spiders of the genus Portia hunt other spiders in ways that seem intelligent, outflanking their victims or luring them from their webs. Laboratory studies show that Portia's instinctive tactics are only starting points for a trial-and-error approach from which these spiders learn very quickly how to overcome new prey species. However, they seem to be relatively slow "thinkers", which is not surprising, as their brains are vastly smaller than those of mammalian predators.

 

Ant-mimicking spiders face several challenges: they generally develop slimmer abdomens and false "waists" in the cephalothorax to mimic the three distinct regions (tagmata) of an ant's body; they wave the first pair of legs in front of their heads to mimic antennae, which spiders lack, and to conceal the fact that they have eight legs rather than six; they develop large color patches round one pair of eyes to disguise the fact that they generally have eight simple eyes, while ants have two compound eyes; they cover their bodies with reflective hairs 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.

 

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 venoms, large jaws or irritant hairs have patches of warning colors, and some actively display these colors when threatened.

 

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 hairs are fine setae (bristles) with fragile bases and a row of barbs on the tip. The barbs cause intense irritation but there is no evidence that they carry any kind of venom. A few defend themselves against wasps by including networks of very robust threads in their webs, giving the spider time to flee while the wasps are struggling with the obstacles. The golden wheeling spider, Carparachne aureoflava, of the Namibian desert escapes parasitic wasps by flipping onto its side and cartwheeling down sand dunes.

 

SOZIAL SPIDERS

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. The genus Anelosimus has a strong tendency towards sociality: all known American species are social, and species in Madagascar are at least somewhat social. Members of other species in the same family but several different genera have independently developed social behavior. For example, although Theridion nigroannulatum belongs to a genus with no other social species, T. nigroannulatum build colonies that may contain several thousand individuals that co-operate in prey capture and share food. Other communal spiders include several Philoponella species (family Uloboridae), Agelena consociata (family Agelenidae) and Mallos gregalis (family Dictynidae). Social predatory spiders need to defend their prey against kleptoparasites ("thieves"), and larger colonies are more successful in this. The herbivorous spider Bagheera kiplingi lives in small colonies which help to protect eggs and spiderlings. Even widow spiders (genus Latrodectus), which are notoriously cannibalistic, have formed small colonies in captivity, sharing webs and feeding together.

 

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 direction required to build them may have been inherited from the common ancestors of most spider groups. However, the majority of spiders build non-orb webs. It used to be thought that the sticky orb web was an evolutionary innovation resulting in the diversification of the Orbiculariae. Now, however, it appears that non-orb spiders are a sub-group 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.

 

ORB WEBS

About half the potential prey that hit orb webs escape. A web has to perform three functions: intercepting the prey (intersection), absorbing its momentum without breaking (stopping), and trapping the prey by entangling it or sticking to it (retention). No single design is best for all prey. For example: wider spacing of lines will increase the web's area and hence its ability to intercept prey, but reduce its stopping power and retention; closer spacing, larger sticky droplets and thicker lines would improve retention, but would make it easier for potential prey to see and avoid the web, at least during the day. However, there are no consistent differences between orb webs built for use during the day and those built for use at night. In fact, there is no simple relationship between orb web design features and the prey they capture, as each orb-weaving species takes a wide range of prey.

 

The hubs of orb webs, where the spiders lurk, are usually above the center, as the spiders can move downwards faster than upwards. If there is an obvious direction in which the spider can retreat to avoid its own predators, the hub is usually offset towards that direction.

 

Horizontal orb webs are fairly common, despite being less effective at intercepting and retaining prey and more vulnerable to damage by rain and falling debris. Various researchers have suggested that horizontal webs offer compensating advantages, such as reduced vulnerability to wind damage; reduced visibility to prey flying upwards, because of the back-lighting 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.

 

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.

 

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.

Tangleweb spiders (cobweb spiders)

 

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.

 

OTHER TYPES OF WEBS

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.

 

EVOLUTION

FOSSIL RECORD

Although the fossil record of spiders is considered poor, almost 1000 species have been described from fossils. Because spiders' bodies are quite soft, the vast majority of fossil spiders have been found preserved in amber. 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 prey, producing silk and possibly caring for their young. In a few cases, amber has preserved spiders' egg sacs and webs, occasionally with prey attached; the oldest fossil web found so far is 100 million years old. Earlier spider fossils come from a few lagerstätten, places where conditions were exceptionally suited to preserving fairly soft tissues.

 

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. 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. 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. 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. Its body length is almost 25 mm.

 

Several Carboniferous spiders were members of the Mesothelae, a primitive group now represented only by the Liphistiidae. The mesothelid Paleothele montceauensis, from the Late Carboniferous over 299 million years ago, had five spinnerets. 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.

 

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.

 

FAMILY TREE

It is now agreed that spiders (Araneae) are monophyletic (i.e., members of a group of organisms that form a clade, consisting of a last common ancestor and all of its descendants). There has been debate about what their closest evolutionary relatives are, and how all of these evolved from the ancestral chelicerates, which were marine animals. The cladogram on the right is based on J. W. Shultz' analysis (2007). Other views include proposals that: scorpions are more closely related to the extinct marine scorpion-like eurypterids than to spiders; spiders and Amblypygi are a monophyletic group. The appearance of several multi-way branchings in the tree on the right shows that there are still uncertainties about relationships between the groups involved.

 

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

 

TAXONOMY

Spiders are divided into two suborders, Mesothelae and Opisthothelae, of which the latter contains two infraorders, Mygalomorphae and Araneomorphae. Nearly 46,000 living species of spiders (order Araneae) have been identified and are currently grouped into about 114 families and about 4,000 genera by arachnologists.

 

WIKIPEDIA

Spiders (order Araneae) are air-breathing arthropods that have eight legs and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other orders of organisms. Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every habitat with the exceptions of air and sea colonization. As of November 2015, at least 45,700 spider species, and 114 families have been recorded by taxonomists. 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.

 

Anatomically, spiders differ from other arthropods in that the usual body segments are fused into two tagmata, the cephalothorax and abdomen, and joined by a small, cylindrical pedicel. Unlike insects, spiders do not have antennae. In all except the most primitive group, the Mesothelae, spiders have the most centralized nervous systems of all arthropods, as all their ganglia are fused into one mass in the cephalothorax. Unlike most arthropods, spiders have no extensor muscles in their limbs and instead extend them by hydraulic pressure.

 

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

 

A herbivorous species, Bagheera kiplingi, was described in 2008, but all other known species are predators, mostly preying on insects and on other spiders, although a few large species also take birds and lizards. Spiders use a wide range of strategies to capture prey: trapping it in sticky webs, lassoing it with sticky bolas, mimicking the prey to avoid detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have acute vision, and hunters of the genus Portia show signs of intelligence in their choice of tactics and ability to develop new ones. Spiders' guts are too narrow to take solids, and they liquefy their food by flooding it with digestive enzymes and grinding it with the bases of their pedipalps, as they do not have true jaws.

 

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

 

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

 

DESCRIPTION

BODY PLAN

Spiders are chelicerates and therefore arthropods. As arthropods they have: segmented bodies with jointed limbs, all covered in a cuticle made of chitin and proteins; heads that are composed of several segments that fuse during the development of the embryo.[7] Being chelicerates, their bodies consist of two tagmata, sets of segments that serve similar functions: the foremost one, called the cephalothorax or prosoma, is a complete fusion of the segments that in an insect would form two separate tagmata, the head and thorax; the rear tagma is called the abdomen or opisthosoma. In spiders, the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel. The pattern of segment fusion that forms chelicerates' heads is unique among arthropods, and what would normally be the first head segment disappears at an early stage of development, so that chelicerates lack the antennae typical of most arthropods. In fact, chelicerates' only appendages ahead of the mouth are a pair of chelicerae, and they lack anything that would function directly as "jaws". The first appendages behind the mouth are called pedipalps, and serve different functions within different groups of chelicerates.

 

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

 

In spiders, the cephalothorax and abdomen are joined by a small, cylindrical pedicel, which enables the abdomen to move independently when producing silk. The upper surface of the cephalothorax is covered by a single, convex 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.

 

Like other arthropods, spiders are coelomates in which the coelom is reduced to small areas round the reproductive and excretory systems. Its place is largely taken by a hemocoel, a cavity that runs most of the length of the body and through which blood flows. The heart is a tube in the upper part of the body, with a few ostia that act as non-return valves allowing blood to enter the heart from the hemocoel but prevent it from leaving before it reaches the front end. 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. The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make oxygen transport more efficient.

 

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. The trachea system has most likely evolved in small ancestors to help resist desiccation. The trachea were originally connected to the surroundings through a pair of openings called spiracles, but in the majority of spiders this pair of spiracles has fused into a single one in the middle, and moved backwards close to the spinnerets. Spiders that have tracheae generally have higher metabolic rates and better water conservation. Spiders are ectotherms, so environmental temperatures affect their activity.

 

FEEDING, DIGESTION AND EXCRETION

Uniquely among chelicerates, the final sections of spiders' chelicerae are fangs, and the great majority of spiders can use them to inject venom into prey from venom glands in the roots of the chelicerae. The family Uloboridae has lost its venom glands, and kills its prey with silk instead. Like most arachnids, including scorpions, spiders have a narrow gut that can only cope with liquid food and spiders have two sets of filters to keep solids out. 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.

 

The stomach in the cephalothorax acts as a pump that sends the food deeper into the digestive system. The mid gut bears many digestive ceca, compartments with no other exit, that extract nutrients from the food; most are in the abdomen, which is dominated by the digestive system, but a few are found in the cephalothorax.

 

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. Production of uric acid and its removal via Malphigian tubules are a water-conserving feature that has evolved independently in several arthropod lineages that can live far away from water,[14] for example the tubules of insects and arachnids develop from completely different parts of the embryo. However, a few primitive spiders, the sub-order Mesothelae and infra-order Mygalomorphae, retain the ancestral arthropod nephridia ("little kidneys"), which use large amounts of water to excrete nitrogenous waste products as ammonia.

 

CENTRAL NERVOS 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. 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; in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfused.

 

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

 

SENSE ORGANS

EYES

Most spiders have four pairs of eyes on the top-front area of the cephalothorax, arranged in patterns that vary from one family to another. The 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, the main eyes at the front of spiders' heads are pigment-cup ocelli that are capable of forming images. The other eyes are thought to be derived from the compound eyes of the ancestral chelicerates, but no longer have the separate facets typical of compound eyes. Unlike the main eyes, in many spiders these secondary eyes detect light reflected from a reflective tapetum lucidum, and wolf spiders can be spotted by torch light reflected from the tapeta. On the other hand, jumping spiders' secondary eyes have no tapeta. Some jumping spiders' visual acuity exceeds by a factor of ten that of dragonflies, which have by far the best vision among insects; in fact the human eye is only about five times sharper than a jumping spider's. They achieve this by a telephoto-like series of lenses, a four-layer retina and the ability to swivel their 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, of these those with six-eyes are the most numerous and are missing a pair of eyes on the anterior median line, others species have four-eyes and some just two. Cave dwelling species have no eyes, or possess vestigial eyes incapable of sight.

 

OTHER SENSES

As with other arthropods, spiders' cuticles would block out information about the outside world, except that they are penetrated by many sensors or connections from sensors to the nervous system. In fact, spiders and other arthropods have modified their cuticles into elaborate arrays of sensors. Various touch sensors, mostly bristles called setae, respond to different levels of force, from strong contact to very weak air currents. Chemical sensors provide equivalents of taste and smell, often by means of setae. Pedipalps carry a large number of such setae sensitive to contact chemicals and air-borne smells, such as female pheromones. Spiders also have in the joints of their limbs slit sensillae that detect forces and vibrations. In web-building spiders, all these mechanical and chemical sensors are more important than the eyes, while the eyes are most important to spiders that hunt actively.

 

Like most arthropods, spiders lack balance and acceleration sensors and rely on their eyes to tell them which way is up. Arthropods' 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.

 

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 exoskeleton to flex their limbs, spiders and a few other groups still use hydraulic pressure to extend them, a system inherited from their pre-arthropod ancestors. The only extensor muscles in spider legs are located in the three hip joints (bordering the coxa and the trochanter). As a result, a spider with a punctured cephalothorax cannot extend its legs, and the legs of dead spiders curl up. Spiders can generate pressures up to eight times their resting level to extend their legs, and jumping spiders can jump up to 50 times their own length by suddenly increasing the blood pressure in the third or fourth pair of legs. Although larger spiders use hydraulics to straighten their legs, unlike smaller jumping spiders they depend on their flexor muscles to generate the propulsive force for their jumps.

 

Most spiders that hunt actively, rather than relying on webs, have dense tufts of fine hairs 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 arachnids, keep at least four legs on the surface while walking or running.

 

SILK PRODUCTION

The abdomen has no appendages except those that have been modified to form one to four (usually three) pairs of short, movable spinnerets, which emit silk. Each spinneret has many spigots, each of which is connected to one silk gland. There are at least six types of silk gland, each producing a different type of silk.

 

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 comb-like 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.

 

Tarantulas also have silk glands in their feet.

 

Even species that do not build webs to catch prey use silk in several ways: as wrappers for sperm and for fertilized eggs; as a "safety rope"; for nest-building; and as "parachutes" by the young of some species.

 

REPRODUCTION AND LIFE CYCLE

Spiders reproduce sexually and fertilization is internal but indirect, in other words the sperm is not inserted into the female's body by the male's genitals but by an intermediate stage. Unlike many land-living arthropods, male spiders do not produce ready-made spermatophores (packages of sperm), but spin small sperm webs on to which they ejaculate and then transfer the sperm to special syringe-like 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".

 

Spiders generally use elaborate courtship rituals to prevent the large females from eating the small males before fertilization, except where the male is so much smaller that he is not worth eating. In web-weaving species, precise patterns of vibrations in the web are a major part of the rituals, while patterns of touches on the female's body are important in many spiders that hunt actively, and may "hypnotize" the female. Gestures and dances by the male are important for jumping spiders, which have excellent eyesight. If courtship is successful, the male injects his sperm from the palpal bulbs into the female's genital opening, known as the epigyne, on the underside of her abdomen. Female's reproductive tracts vary from simple tubes to systems that include seminal receptacles in which females store sperm and release it when they are ready.

 

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 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 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 eggs in one or more silk egg sacs, which maintain a fairly constant humidity level. In some species, the females die afterwards, but females of other species protect the sacs by attaching them to their webs, hiding them in nests, carrying them in the chelicerae or attaching them to the spinnerets and dragging them along.

 

Baby spiders pass all their larval stages inside the egg and hatch 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 cling to rough bristles on the mother's back, and females of some species respond to the "begging" behaviour of their young by giving them their prey, provided it is no longer struggling, or even regurgitate food.

 

Like other arthropods, spiders have to molt to grow as their cuticle ("skin") cannot stretch. In some species males mate with newly molted females, which are too weak to be dangerous to the males. Most spiders live for only one to two years, although some tarantulas can live in captivity for over 20 years.

 

SIZE

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

 

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.

 

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 amino acids, lipids, vitamins and minerals in addition to sugars, and studies have shown that other spider species live longer when nectar is available. Feeding on nectar avoids the risks of struggles with prey, and the costs of producing venom and digestive enzymes.

 

Various species are known to feed on dead arthropods (scavenging), web silk, and their own shed exoskeletons. Pollen caught in webs may also be eaten, and studies have shown that young spiders have a better chance of survival if they have the opportunity to eat pollen. In captivity, several spider species are also known to feed on bananas, marmalade, milk, egg yolk and sausages.

 

METHODS OF CAPTURING PREY

The best-known method of prey capture is by means of sticky webs. Varying placement of webs allows different species of spider to trap different insects in the same area, for example flat horizontal webs trap insects that fly up from vegetation underneath while flat vertical webs trap insects in horizontal flight. Web-building spiders have poor vision, but are extremely sensitive to vibrations.

 

Females of the water spider Argyroneta aquatica build underwater "diving bell" webs that they fill with air and use for digesting prey, molting, mating and raising offspring. They live almost entirely within the bells, darting out to catch prey animals that touch the bell or the threads that anchor it. A few spiders use the surfaces of lakes and ponds as "webs", detecting trapped insects by the vibrations that these cause while struggling.

 

Net-casting spiders weave only small webs, but then manipulate them to trap prey. Those of the genus Hyptiotes 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 tipulid flies that they catch with the backwards strike.

 

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. Juveniles and adult males are much smaller and do not make bolas. Instead they release different pheromones that attract moth flies, and catch them with their front pairs of legs.

 

The primitive Liphistiidae, the "trapdoor spiders" of the family Ctenizidae and many tarantulas are ambush 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. Other ambush predators do without such aids, including many crab spiders, and a few species that prey on bees, which see ultraviolet, can adjust their ultraviolet reflectance to match the flowers in which they are lurking. Wolf spiders, jumping spiders, fishing spiders and some crab spiders capture prey by chasing it, and rely mainly on vision to locate prey.Some jumping spiders of the genus Portia hunt other spiders in ways that seem intelligent, outflanking their victims or luring them from their webs. Laboratory studies show that Portia's instinctive tactics are only starting points for a trial-and-error approach from which these spiders learn very quickly how to overcome new prey species. However, they seem to be relatively slow "thinkers", which is not surprising, as their brains are vastly smaller than those of mammalian predators.

 

Ant-mimicking spiders face several challenges: they generally develop slimmer abdomens and false "waists" in the cephalothorax to mimic the three distinct regions (tagmata) of an ant's body; they wave the first pair of legs in front of their heads to mimic antennae, which spiders lack, and to conceal the fact that they have eight legs rather than six; they develop large color patches round one pair of eyes to disguise the fact that they generally have eight simple eyes, while ants have two compound eyes; they cover their bodies with reflective hairs 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.

 

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 venoms, large jaws or irritant hairs have patches of warning colors, and some actively display these colors when threatened.

 

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 hairs are fine setae (bristles) with fragile bases and a row of barbs on the tip. The barbs cause intense irritation but there is no evidence that they carry any kind of venom. A few defend themselves against wasps by including networks of very robust threads in their webs, giving the spider time to flee while the wasps are struggling with the obstacles. The golden wheeling spider, Carparachne aureoflava, of the Namibian desert escapes parasitic wasps by flipping onto its side and cartwheeling down sand dunes.

 

SOZIAL SPIDERS

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. The genus Anelosimus has a strong tendency towards sociality: all known American species are social, and species in Madagascar are at least somewhat social. Members of other species in the same family but several different genera have independently developed social behavior. For example, although Theridion nigroannulatum belongs to a genus with no other social species, T. nigroannulatum build colonies that may contain several thousand individuals that co-operate in prey capture and share food. Other communal spiders include several Philoponella species (family Uloboridae), Agelena consociata (family Agelenidae) and Mallos gregalis (family Dictynidae). Social predatory spiders need to defend their prey against kleptoparasites ("thieves"), and larger colonies are more successful in this. The herbivorous spider Bagheera kiplingi lives in small colonies which help to protect eggs and spiderlings. Even widow spiders (genus Latrodectus), which are notoriously cannibalistic, have formed small colonies in captivity, sharing webs and feeding together.

 

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 direction required to build them may have been inherited from the common ancestors of most spider groups. However, the majority of spiders build non-orb webs. It used to be thought that the sticky orb web was an evolutionary innovation resulting in the diversification of the Orbiculariae. Now, however, it appears that non-orb spiders are a sub-group 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.

 

ORB WEBS

About half the potential prey that hit orb webs escape. A web has to perform three functions: intercepting the prey (intersection), absorbing its momentum without breaking (stopping), and trapping the prey by entangling it or sticking to it (retention). No single design is best for all prey. For example: wider spacing of lines will increase the web's area and hence its ability to intercept prey, but reduce its stopping power and retention; closer spacing, larger sticky droplets and thicker lines would improve retention, but would make it easier for potential prey to see and avoid the web, at least during the day. However, there are no consistent differences between orb webs built for use during the day and those built for use at night. In fact, there is no simple relationship between orb web design features and the prey they capture, as each orb-weaving species takes a wide range of prey.

 

The hubs of orb webs, where the spiders lurk, are usually above the center, as the spiders can move downwards faster than upwards. If there is an obvious direction in which the spider can retreat to avoid its own predators, the hub is usually offset towards that direction.

 

Horizontal orb webs are fairly common, despite being less effective at intercepting and retaining prey and more vulnerable to damage by rain and falling debris. Various researchers have suggested that horizontal webs offer compensating advantages, such as reduced vulnerability to wind damage; reduced visibility to prey flying upwards, because of the back-lighting 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.

 

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.

 

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.

Tangleweb spiders (cobweb spiders)

 

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.

 

OTHER TYPES OF WEBS

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.

 

EVOLUTION

FOSSIL RECORD

Although the fossil record of spiders is considered poor, almost 1000 species have been described from fossils. Because spiders' bodies are quite soft, the vast majority of fossil spiders have been found preserved in amber. 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 prey, producing silk and possibly caring for their young. In a few cases, amber has preserved spiders' egg sacs and webs, occasionally with prey attached; the oldest fossil web found so far is 100 million years old. Earlier spider fossils come from a few lagerstätten, places where conditions were exceptionally suited to preserving fairly soft tissues.

 

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. 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. 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. 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. Its body length is almost 25 mm.

 

Several Carboniferous spiders were members of the Mesothelae, a primitive group now represented only by the Liphistiidae. The mesothelid Paleothele montceauensis, from the Late Carboniferous over 299 million years ago, had five spinnerets. 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.

 

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.

 

FAMILY TREE

It is now agreed that spiders (Araneae) are monophyletic (i.e., members of a group of organisms that form a clade, consisting of a last common ancestor and all of its descendants). There has been debate about what their closest evolutionary relatives are, and how all of these evolved from the ancestral chelicerates, which were marine animals. The cladogram on the right is based on J. W. Shultz' analysis (2007). Other views include proposals that: scorpions are more closely related to the extinct marine scorpion-like eurypterids than to spiders; spiders and Amblypygi are a monophyletic group. The appearance of several multi-way branchings in the tree on the right shows that there are still uncertainties about relationships between the groups involved.

 

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

 

TAXONOMY

Spiders are divided into two suborders, Mesothelae and Opisthothelae, of which the latter contains two infraorders, Mygalomorphae and Araneomorphae. Nearly 46,000 living species of spiders (order Araneae) have been identified and are currently grouped into about 114 families and about 4,000 genera by arachnologists.

 

WIKIPEDIA

Spiders (order Araneae) are air-breathing arthropods that have eight legs and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other orders of organisms. Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every habitat with the exceptions of air and sea colonization. As of November 2015, at least 45,700 spider species, and 114 families have been recorded by taxonomists. 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.

 

Anatomically, spiders differ from other arthropods in that the usual body segments are fused into two tagmata, the cephalothorax and abdomen, and joined by a small, cylindrical pedicel. Unlike insects, spiders do not have antennae. In all except the most primitive group, the Mesothelae, spiders have the most centralized nervous systems of all arthropods, as all their ganglia are fused into one mass in the cephalothorax. Unlike most arthropods, spiders have no extensor muscles in their limbs and instead extend them by hydraulic pressure.

 

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

 

A herbivorous species, Bagheera kiplingi, was described in 2008,[5] but all other known species are predators, mostly preying on insects and on other spiders, although a few large species also take birds and lizards. Spiders use a wide range of strategies to capture prey: trapping it in sticky webs, lassoing it with sticky bolas, mimicking the prey to avoid detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have acute vision, and hunters of the genus Portia show signs of intelligence in their choice of tactics and ability to develop new ones. Spiders' guts are too narrow to take solids, and they liquefy their food by flooding it with digestive enzymes and grinding it with the bases of their pedipalps, as they do not have true jaws.

 

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

 

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

 

BODY PLAN

Spiders are chelicerates and therefore arthropods.[6] As arthropods they have: segmented bodies with jointed limbs, all covered in a cuticle made of chitin and proteins; heads that are composed of several segments that fuse during the development of the embryo. 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. In spiders, the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel. The pattern of segment fusion that forms chelicerates' heads is unique among arthropods, and what would normally be the first head segment disappears at an early stage of development, so that chelicerates lack the antennae typical of most arthropods. In fact, chelicerates' only appendages ahead of the mouth are a pair of chelicerae, and they lack anything that would function directly as "jaws". The first appendages behind the mouth are called pedipalps, and serve different functions within different groups of chelicerates.

 

Spiders and scorpions are members of one chelicerate group, the arachnids. Scorpions' chelicerae have three sections and are used in feeding. Spiders' chelicerae have two sections and terminate in fangs that are generally venomous, and fold away behind the upper sections while not in use. The upper sections generally have thick "beards" that filter solid lumps out of their food, as spiders can take only liquid food.[8] Scorpions' pedipalps generally form large claws for capturing prey, while those of spiders are fairly small appendages whose bases also act as an extension of the mouth; in addition, those of male spiders have enlarged last sections used for sperm transfer.

 

In spiders, the cephalothorax and abdomen are joined by a small, cylindrical pedicel, which enables the abdomen to move independently when producing silk. The upper surface of the cephalothorax is covered by a single, convex 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.

 

CIRCULATION AND RESPIRATION

Like other arthropods, spiders are coelomates in which the coelom is reduced to small areas round the reproductive and excretory systems. Its place is largely taken by a hemocoel, a cavity that runs most of the length of the body and through which blood flows. The heart is a tube in the upper part of the body, with a few ostia that act as non-return valves allowing blood to enter the heart from the hemocoel but prevent it from leaving before it reaches the front end. 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. The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make oxygen transport more efficient.

 

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. The trachea system has most likely evolved in small ancestors to help resist desiccation. The trachea were originally connected to the surroundings through a pair of openings called spiracles, but in the majority of spiders this pair of spiracles has fused into a single one in the middle, and moved backwards close to the spinnerets. Spiders that have tracheae generally have higher metabolic rates and better water conservation. Spiders are ectotherms, so environmental temperatures affect their activity.

 

FEEDING, DIGESTION AND EXCRETION

Uniquely among chelicerates, the final sections of spiders' chelicerae are fangs, and the great majority of spiders can use them to inject venom into prey from venom glands in the roots of the chelicerae. The family Uloboridae has lost its venom glands, and kills its prey with silk instead. Like most arachnids, including scorpions, spiders have a narrow gut that can only cope with liquid food and spiders have two sets of filters to keep solids out. 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.

 

The stomach in the cephalothorax acts as a pump that sends the food deeper into the digestive system. The mid gut bears many digestive ceca, compartments with no other exit, that extract nutrients from the food; most are in the abdomen, which is dominated by the digestive system, but a few are found in the cephalothorax.

 

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. 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 insects and arachnids develop from completely different parts of the embryo. However, a few primitive spiders, the sub-order Mesothelae and infra-order Mygalomorphae, retain the ancestral arthropod nephridia ("little kidneys"), which use large amounts of water to excrete nitrogenous waste products as ammonia.

 

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. 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; in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfused.

 

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

Sense organs

 

EYES

Most spiders have four pairs of eyes on the top-front area of the cephalothorax, arranged in patterns that vary from one family to another. The 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, the main eyes at the front of spiders' heads are pigment-cup ocelli that are capable of forming images. The other eyes are thought to be derived from the compound eyes of the ancestral chelicerates, but no longer have the separate facets typical of compound eyes. Unlike the main eyes, in many spiders these secondary eyes detect light reflected from a reflective tapetum lucidum, and wolf spiders can be spotted by torch light reflected from the tapeta. On the other hand, jumping spiders' secondary eyes have no tapeta. Some jumping spiders' visual acuity exceeds by a factor of ten that of dragonflies, which have by far the best vision among insects; in fact the human eye is only about five times sharper than a jumping spider's. They achieve this by a telephoto-like series of lenses, a four-layer retina and the ability to swivel their 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, of these those with six-eyes are the most numerous and are missing a pair of eyes on the anterior median line, others species have four-eyes and some just two. Cave dwelling species have no eyes, or possess vestigial eyes incapable of sight.

 

OTHER SENSES

As with other arthropods, spiders' cuticles would block out information about the outside world, except that they are penetrated by many sensors or connections from sensors to the nervous system. In fact, spiders and other arthropods have modified their cuticles into elaborate arrays of sensors. Various touch sensors, mostly bristles called setae, respond to different levels of force, from strong contact to very weak air currents. Chemical sensors provide equivalents of taste and smell, often by means of setae. Pedipalps carry a large number of such setae sensitive to contact chemicals and air-borne smells, such as female pheromones. Spiders also have in the joints of their limbs slit sensillae that detect forces and vibrations. In web-building spiders, all these mechanical and chemical sensors are more important than the eyes, while the eyes are most important to spiders that hunt actively.

 

Like most arthropods, spiders lack balance and acceleration sensors and rely on their eyes to tell them which way is up. Arthropods' 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.

 

LOCMOTION

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 exoskeleton to flex their limbs, spiders and a few other groups still use hydraulic pressure to extend them, a system inherited from their pre-arthropod ancestors. The only extensor muscles in spider legs are located in the three hip joints (bordering the coxa and the trochanter). As a result, a spider with a punctured cephalothorax cannot extend its legs, and the legs of dead spiders curl up. Spiders can generate pressures up to eight times their resting level to extend their legs, and jumping spiders can jump up to 50 times their own length by suddenly increasing the blood pressure in the third or fourth pair of legs. Although larger spiders use hydraulics to straighten their legs, unlike smaller jumping spiders they depend on their flexor muscles to generate the propulsive force for their jumps.

 

Most spiders that hunt actively, rather than relying on webs, have dense tufts of fine hairs 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.[8] Spiders, like most other arachnids, keep at least four legs on the surface while walking or running.

 

SILK PRODUCTION

The abdomen has no appendages except those that have been modified to form one to four (usually three) pairs of short, movable spinnerets, which emit silk. Each spinneret has many spigots, each of which is connected to one silk gland. There are at least six types of silk gland, each producing a different type of silk.

 

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 comb-like 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.

 

Tarantulas also have silk glands in their feet.

 

Even species that do not build webs to catch prey use silk in several ways: as wrappers for sperm and for fertilized eggs; as a "safety rope"; for nest-building; and as "parachutes" by the young of some species.

 

REPRODUCTION AND LIFE CYCLE

Spiders reproduce sexually and fertilization is internal but indirect, in other words the sperm is not inserted into the female's body by the male's genitals but by an intermediate stage. Unlike many land-living arthropods, male spiders do not produce ready-made spermatophores (packages of sperm), but spin small sperm webs on to which they ejaculate and then transfer the sperm to special syringe-like 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".

 

Spiders generally use elaborate courtship rituals to prevent the large females from eating the small males before fertilization, except where the male is so much smaller that he is not worth eating. In web-weaving species, precise patterns of vibrations in the web are a major part of the rituals, while patterns of touches on the female's body are important in many spiders that hunt actively, and may "hypnotize" the female. Gestures and dances by the male are important for jumping spiders, which have excellent eyesight. If courtship is successful, the male injects his sperm from the palpal bulbs into the female's genital opening, known as the epigyne, on the underside of her abdomen. Female's reproductive tracts vary from simple tubes to systems that include seminal receptacles in which females store sperm and release it when they are ready.

 

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 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 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 eggs in one or more silk egg sacs, which maintain a fairly constant humidity level. In some species, the females die afterwards, but females of other species protect the sacs by attaching them to their webs, hiding them in nests, carrying them in the chelicerae or attaching them to the spinnerets and dragging them along.

 

Baby spiders pass all their larval stages inside the egg and hatch 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 cling to rough bristles on the mother's back, and females of some species respond to the "begging" behaviour of their young by giving them their prey, provided it is no longer struggling, or even regurgitate food.

 

Like other arthropods, spiders have to molt to grow as their cuticle ("skin") cannot stretch. In some species males mate with newly molted females, which are too weak to be dangerous to the males. Most spiders live for only one to two years, although some tarantulas can live in captivity for over 20 years.

 

SIZE

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

 

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 brown coloration. Bilins are found, for example, in Micrommata virescens, resulting in its green color. Guanine is responsible for the white markings of the European garden spider Araneus diadematus. 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. 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 hairs reflecting the light, Lycosa and Josa both have areas of modified cuticle that act as light reflectors.

 

ECOGOGY 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.

 

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 amino acids, lipids, vitamins and minerals in addition to sugars, and studies have shown that other spider species live longer when nectar is available. Feeding on nectar avoids the risks of struggles with prey, and the costs of producing venom and digestive enzymes.

 

Various species are known to feed on dead arthropods (scavenging), web silk, and their own shed exoskeletons. Pollen caught in webs may also be eaten, and studies have shown that young spiders have a better chance of survival if they have the opportunity to eat pollen. In captivity, several spider species are also known to feed on bananas, marmalade, milk, egg yolk and sausages.

 

METHODS OF CAPTURING PREY

The best-known method of prey capture is by means of sticky webs. Varying placement of webs allows different species of spider to trap different insects in the same area, for example flat horizontal webs trap insects that fly up from vegetation underneath while flat vertical webs trap insects in horizontal flight. Web-building spiders have poor vision, but are extremely sensitive to vibrations.

 

Females of the water spider Argyroneta aquatica build underwater "diving bell" webs that they fill with air and use for digesting prey, molting, mating and raising offspring. They live almost entirely within the bells, darting out to catch prey animals that touch the bell or the threads that anchor it. A few spiders use the surfaces of lakes and ponds as "webs", detecting trapped insects by the vibrations that these cause while struggling.

 

Net-casting spiders weave only small webs, but then manipulate them to trap prey. Those of the genus Hyptiotes 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 tipulid flies that they catch with the backwards strike.

 

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. Juveniles and adult males are much smaller and do not make bolas. Instead they release different pheromones that attract moth flies, and catch them with their front pairs of legs.

 

The primitive Liphistiidae, the "trapdoor spiders" of the family Ctenizidae and many tarantulas are ambush 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. Other ambush predators do without such aids, including many crab spiders, and a few species that prey on bees, which see ultraviolet, can adjust their ultraviolet reflectance to match the flowers in which they are lurking. Wolf spiders, jumping spiders, fishing spiders and some crab spiders capture prey by chasing it, and rely mainly on vision to locate prey.Some jumping spiders of the genus Portia hunt other spiders in ways that seem intelligent, outflanking their victims or luring them from their webs. Laboratory studies show that Portia's instinctive tactics are only starting points for a trial-and-error approach from which these spiders learn very quickly how to overcome new prey species. However, they seem to be relatively slow "thinkers", which is not surprising, as their brains are vastly smaller than those of mammalian predators.Ant-mimicking spiders face several challenges: they generally develop slimmer abdomens and false "waists" in the cephalothorax to mimic the three distinct regions (tagmata) of an ant's body; they wave the first pair of legs in front of their heads to mimic antennae, which spiders lack, and to conceal the fact that they have eight legs rather than six; they develop large color patches round one pair of eyes to disguise the fact that they generally have eight simple eyes, while ants have two compound eyes; they cover their bodies with reflective hairs 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.

 

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 venoms, large jaws or irritant hairs have patches of warning colors, and some actively display these colors when threatened.

 

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 hairs are fine setae (bristles) with fragile bases and a row of barbs on the tip. The barbs cause intense irritation but there is no evidence that they carry any kind of venom. A few defend themselves against wasps by including networks of very robust threads in their webs, giving the spider time to flee while the wasps are struggling with the obstacles. The golden wheeling spider, Carparachne aureoflava, of the Namibian desert escapes parasitic wasps by flipping onto its side and cartwheeling down sand dunes.

 

SOCIAL SPIDERS

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. The genus Anelosimus has a strong tendency towards sociality: all known American species are social, and species in Madagascar are at least somewhat social. Members of other species in the same family but several different genera have independently developed social behavior. For example, although Theridion nigroannulatum belongs to a genus with no other social species, T. nigroannulatum build colonies that may contain several thousand individuals that co-operate in prey capture and share food. Other communal spiders include several Philoponella species (family Uloboridae), Agelena consociata (family Agelenidae) and Mallos gregalis (family Dictynidae). Social predatory spiders need to defend their prey against kleptoparasites ("thieves"), and larger colonies are more successful in this. The herbivorous spider Bagheera kiplingi lives in small colonies which help to protect eggs and spiderlings. Even widow spiders (genus Latrodectus), which are notoriously cannibalistic, have formed small colonies in captivity, sharing webs and feeding together.

 

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 direction required to build them may have been inherited from the common ancestors of most spider groups. However, the majority of spiders build non-orb webs. It used to be thought that the sticky orb web was an evolutionary innovation resulting in the diversification of the Orbiculariae. Now, however, it appears that non-orb spiders are a sub-group 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.

 

ORB WEBS

About half the potential prey that hit orb webs escape. A web has to perform three functions: intercepting the prey (intersection), absorbing its momentum without breaking (stopping), and trapping the prey by entangling it or sticking to it (retention). No single design is best for all prey. For example: wider spacing of lines will increase the web's area and hence its ability to intercept prey, but reduce its stopping power and retention; closer spacing, larger sticky droplets and thicker lines would improve retention, but would make it easier for potential prey to see and avoid the web, at least during the day. However, there are no consistent differences between orb webs built for use during the day and those built for use at night. In fact, there is no simple relationship between orb web design features and the prey they capture, as each orb-weaving species takes a wide range of prey.

 

The hubs of orb webs, where the spiders lurk, are usually above the center, as the spiders can move downwards faster than upwards. If there is an obvious direction in which the spider can retreat to avoid its own predators, the hub is usually offset towards that direction.

 

Horizontal orb webs are fairly common, despite being less effective at intercepting and retaining prey and more vulnerable to damage by rain and falling debris. Various researchers have suggested that horizontal webs offer compensating advantages, such as reduced vulnerability to wind damage; reduced visibility to prey flying upwards, because of the back-lighting 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.

 

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; "ladder-like" webs that appear most effective in catching moths. However, the significance of many variations is unclear.

 

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.

 

TANGLEWEB SPIDERS (COBWEB SPIDERS)

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.

 

OTHER TYPES OF WEBS

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.

 

EVOLUTION

FOSSIL RECORD

Although the fossil record of spiders is considered poor, almost 1000 species have been described from fossils. Because spiders' bodies are quite soft, the vast majority of fossil spiders have been found preserved in amber. 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 prey, producing silk and possibly caring for their young. In a few cases, amber has preserved spiders' egg sacs and webs, occasionally with prey attached; the oldest fossil web found so far is 100 million years old. Earlier spider fossils come from a few lagerstätten, places where conditions were exceptionally suited to preserving fairly soft tissues.

 

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. 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. 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. 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. Its body length is almost 25 mm.

 

Several Carboniferous spiders were members of the Mesothelae, a primitive group now represented only by the Liphistiidae. The mesothelid Paleothele montceauensis, from the Late Carboniferous over 299 million years ago, had five spinnerets. 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.

 

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.

 

WIKIPEDIA

Spiders (order Araneae) are air-breathing arthropods that have eight legs and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other orders of organisms. Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every habitat with the exceptions of air and sea colonization. As of November 2015, at least 45,700 spider species, and 114 families have been recorded by taxonomists. 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.

 

Anatomically, spiders differ from other arthropods in that the usual body segments are fused into two tagmata, the cephalothorax and abdomen, and joined by a small, cylindrical pedicel. Unlike insects, spiders do not have antennae. In all except the most primitive group, the Mesothelae, spiders have the most centralized nervous systems of all arthropods, as all their ganglia are fused into one mass in the cephalothorax. Unlike most arthropods, spiders have no extensor muscles in their limbs and instead extend them by hydraulic pressure.

 

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

 

A herbivorous species, Bagheera kiplingi, was described in 2008,[5] but all other known species are predators, mostly preying on insects and on other spiders, although a few large species also take birds and lizards. Spiders use a wide range of strategies to capture prey: trapping it in sticky webs, lassoing it with sticky bolas, mimicking the prey to avoid detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have acute vision, and hunters of the genus Portia show signs of intelligence in their choice of tactics and ability to develop new ones. Spiders' guts are too narrow to take solids, and they liquefy their food by flooding it with digestive enzymes and grinding it with the bases of their pedipalps, as they do not have true jaws.

 

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

 

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

 

BODY PLAN

Spiders are chelicerates and therefore arthropods.[6] As arthropods they have: segmented bodies with jointed limbs, all covered in a cuticle made of chitin and proteins; heads that are composed of several segments that fuse during the development of the embryo. 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. In spiders, the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel. The pattern of segment fusion that forms chelicerates' heads is unique among arthropods, and what would normally be the first head segment disappears at an early stage of development, so that chelicerates lack the antennae typical of most arthropods. In fact, chelicerates' only appendages ahead of the mouth are a pair of chelicerae, and they lack anything that would function directly as "jaws". The first appendages behind the mouth are called pedipalps, and serve different functions within different groups of chelicerates.

 

Spiders and scorpions are members of one chelicerate group, the arachnids. Scorpions' chelicerae have three sections and are used in feeding. Spiders' chelicerae have two sections and terminate in fangs that are generally venomous, and fold away behind the upper sections while not in use. The upper sections generally have thick "beards" that filter solid lumps out of their food, as spiders can take only liquid food.[8] Scorpions' pedipalps generally form large claws for capturing prey, while those of spiders are fairly small appendages whose bases also act as an extension of the mouth; in addition, those of male spiders have enlarged last sections used for sperm transfer.

 

In spiders, the cephalothorax and abdomen are joined by a small, cylindrical pedicel, which enables the abdomen to move independently when producing silk. The upper surface of the cephalothorax is covered by a single, convex 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.

 

CIRCULATION AND RESPIRATION

Like other arthropods, spiders are coelomates in which the coelom is reduced to small areas round the reproductive and excretory systems. Its place is largely taken by a hemocoel, a cavity that runs most of the length of the body and through which blood flows. The heart is a tube in the upper part of the body, with a few ostia that act as non-return valves allowing blood to enter the heart from the hemocoel but prevent it from leaving before it reaches the front end. 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. The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make oxygen transport more efficient.

 

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. The trachea system has most likely evolved in small ancestors to help resist desiccation. The trachea were originally connected to the surroundings through a pair of openings called spiracles, but in the majority of spiders this pair of spiracles has fused into a single one in the middle, and moved backwards close to the spinnerets. Spiders that have tracheae generally have higher metabolic rates and better water conservation. Spiders are ectotherms, so environmental temperatures affect their activity.

 

FEEDING, DIGESTION AND EXCRETION

Uniquely among chelicerates, the final sections of spiders' chelicerae are fangs, and the great majority of spiders can use them to inject venom into prey from venom glands in the roots of the chelicerae. The family Uloboridae has lost its venom glands, and kills its prey with silk instead. Like most arachnids, including scorpions, spiders have a narrow gut that can only cope with liquid food and spiders have two sets of filters to keep solids out. 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.

 

The stomach in the cephalothorax acts as a pump that sends the food deeper into the digestive system. The mid gut bears many digestive ceca, compartments with no other exit, that extract nutrients from the food; most are in the abdomen, which is dominated by the digestive system, but a few are found in the cephalothorax.

 

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. 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 insects and arachnids develop from completely different parts of the embryo. However, a few primitive spiders, the sub-order Mesothelae and infra-order Mygalomorphae, retain the ancestral arthropod nephridia ("little kidneys"), which use large amounts of water to excrete nitrogenous waste products as ammonia.

 

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. 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; in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfused.

 

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

Sense organs

 

EYES

Most spiders have four pairs of eyes on the top-front area of the cephalothorax, arranged in patterns that vary from one family to another. The 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, the main eyes at the front of spiders' heads are pigment-cup ocelli that are capable of forming images. The other eyes are thought to be derived from the compound eyes of the ancestral chelicerates, but no longer have the separate facets typical of compound eyes. Unlike the main eyes, in many spiders these secondary eyes detect light reflected from a reflective tapetum lucidum, and wolf spiders can be spotted by torch light reflected from the tapeta. On the other hand, jumping spiders' secondary eyes have no tapeta. Some jumping spiders' visual acuity exceeds by a factor of ten that of dragonflies, which have by far the best vision among insects; in fact the human eye is only about five times sharper than a jumping spider's. They achieve this by a telephoto-like series of lenses, a four-layer retina and the ability to swivel their 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, of these those with six-eyes are the most numerous and are missing a pair of eyes on the anterior median line, others species have four-eyes and some just two. Cave dwelling species have no eyes, or possess vestigial eyes incapable of sight.

 

OTHER SENSES

As with other arthropods, spiders' cuticles would block out information about the outside world, except that they are penetrated by many sensors or connections from sensors to the nervous system. In fact, spiders and other arthropods have modified their cuticles into elaborate arrays of sensors. Various touch sensors, mostly bristles called setae, respond to different levels of force, from strong contact to very weak air currents. Chemical sensors provide equivalents of taste and smell, often by means of setae. Pedipalps carry a large number of such setae sensitive to contact chemicals and air-borne smells, such as female pheromones. Spiders also have in the joints of their limbs slit sensillae that detect forces and vibrations. In web-building spiders, all these mechanical and chemical sensors are more important than the eyes, while the eyes are most important to spiders that hunt actively.

 

Like most arthropods, spiders lack balance and acceleration sensors and rely on their eyes to tell them which way is up. Arthropods' 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.

 

LOCMOTION

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 exoskeleton to flex their limbs, spiders and a few other groups still use hydraulic pressure to extend them, a system inherited from their pre-arthropod ancestors. The only extensor muscles in spider legs are located in the three hip joints (bordering the coxa and the trochanter). As a result, a spider with a punctured cephalothorax cannot extend its legs, and the legs of dead spiders curl up. Spiders can generate pressures up to eight times their resting level to extend their legs, and jumping spiders can jump up to 50 times their own length by suddenly increasing the blood pressure in the third or fourth pair of legs. Although larger spiders use hydraulics to straighten their legs, unlike smaller jumping spiders they depend on their flexor muscles to generate the propulsive force for their jumps.

 

Most spiders that hunt actively, rather than relying on webs, have dense tufts of fine hairs 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.[8] Spiders, like most other arachnids, keep at least four legs on the surface while walking or running.

 

SILK PRODUCTION

The abdomen has no appendages except those that have been modified to form one to four (usually three) pairs of short, movable spinnerets, which emit silk. Each spinneret has many spigots, each of which is connected to one silk gland. There are at least six types of silk gland, each producing a different type of silk.

 

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 comb-like 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.

 

Tarantulas also have silk glands in their feet.

 

Even species that do not build webs to catch prey use silk in several ways: as wrappers for sperm and for fertilized eggs; as a "safety rope"; for nest-building; and as "parachutes" by the young of some species.

 

REPRODUCTION AND LIFE CYCLE

Spiders reproduce sexually and fertilization is internal but indirect, in other words the sperm is not inserted into the female's body by the male's genitals but by an intermediate stage. Unlike many land-living arthropods, male spiders do not produce ready-made spermatophores (packages of sperm), but spin small sperm webs on to which they ejaculate and then transfer the sperm to special syringe-like 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".

 

Spiders generally use elaborate courtship rituals to prevent the large females from eating the small males before fertilization, except where the male is so much smaller that he is not worth eating. In web-weaving species, precise patterns of vibrations in the web are a major part of the rituals, while patterns of touches on the female's body are important in many spiders that hunt actively, and may "hypnotize" the female. Gestures and dances by the male are important for jumping spiders, which have excellent eyesight. If courtship is successful, the male injects his sperm from the palpal bulbs into the female's genital opening, known as the epigyne, on the underside of her abdomen. Female's reproductive tracts vary from simple tubes to systems that include seminal receptacles in which females store sperm and release it when they are ready.

 

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 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 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 eggs in one or more silk egg sacs, which maintain a fairly constant humidity level. In some species, the females die afterwards, but females of other species protect the sacs by attaching them to their webs, hiding them in nests, carrying them in the chelicerae or attaching them to the spinnerets and dragging them along.

 

Baby spiders pass all their larval stages inside the egg and hatch 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 cling to rough bristles on the mother's back, and females of some species respond to the "begging" behaviour of their young by giving them their prey, provided it is no longer struggling, or even regurgitate food.

 

Like other arthropods, spiders have to molt to grow as their cuticle ("skin") cannot stretch. In some species males mate with newly molted females, which are too weak to be dangerous to the males. Most spiders live for only one to two years, although some tarantulas can live in captivity for over 20 years.

 

SIZE

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

 

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 brown coloration. Bilins are found, for example, in Micrommata virescens, resulting in its green color. Guanine is responsible for the white markings of the European garden spider Araneus diadematus. 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. 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 hairs reflecting the light, Lycosa and Josa both have areas of modified cuticle that act as light reflectors.

 

ECOGOGY 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.

 

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 amino acids, lipids, vitamins and minerals in addition to sugars, and studies have shown that other spider species live longer when nectar is available. Feeding on nectar avoids the risks of struggles with prey, and the costs of producing venom and digestive enzymes.

 

Various species are known to feed on dead arthropods (scavenging), web silk, and their own shed exoskeletons. Pollen caught in webs may also be eaten, and studies have shown that young spiders have a better chance of survival if they have the opportunity to eat pollen. In captivity, several spider species are also known to feed on bananas, marmalade, milk, egg yolk and sausages.

 

METHODS OF CAPTURING PREY

The best-known method of prey capture is by means of sticky webs. Varying placement of webs allows different species of spider to trap different insects in the same area, for example flat horizontal webs trap insects that fly up from vegetation underneath while flat vertical webs trap insects in horizontal flight. Web-building spiders have poor vision, but are extremely sensitive to vibrations.

 

Females of the water spider Argyroneta aquatica build underwater "diving bell" webs that they fill with air and use for digesting prey, molting, mating and raising offspring. They live almost entirely within the bells, darting out to catch prey animals that touch the bell or the threads that anchor it. A few spiders use the surfaces of lakes and ponds as "webs", detecting trapped insects by the vibrations that these cause while struggling.

 

Net-casting spiders weave only small webs, but then manipulate them to trap prey. Those of the genus Hyptiotes 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 tipulid flies that they catch with the backwards strike.

 

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. Juveniles and adult males are much smaller and do not make bolas. Instead they release different pheromones that attract moth flies, and catch them with their front pairs of legs.

 

The primitive Liphistiidae, the "trapdoor spiders" of the family Ctenizidae and many tarantulas are ambush 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. Other ambush predators do without such aids, including many crab spiders, and a few species that prey on bees, which see ultraviolet, can adjust their ultraviolet reflectance to match the flowers in which they are lurking. Wolf spiders, jumping spiders, fishing spiders and some crab spiders capture prey by chasing it, and rely mainly on vision to locate prey.Some jumping spiders of the genus Portia hunt other spiders in ways that seem intelligent, outflanking their victims or luring them from their webs. Laboratory studies show that Portia's instinctive tactics are only starting points for a trial-and-error approach from which these spiders learn very quickly how to overcome new prey species. However, they seem to be relatively slow "thinkers", which is not surprising, as their brains are vastly smaller than those of mammalian predators.Ant-mimicking spiders face several challenges: they generally develop slimmer abdomens and false "waists" in the cephalothorax to mimic the three distinct regions (tagmata) of an ant's body; they wave the first pair of legs in front of their heads to mimic antennae, which spiders lack, and to conceal the fact that they have eight legs rather than six; they develop large color patches round one pair of eyes to disguise the fact that they generally have eight simple eyes, while ants have two compound eyes; they cover their bodies with reflective hairs 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.

 

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 venoms, large jaws or irritant hairs have patches of warning colors, and some actively display these colors when threatened.

 

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 hairs are fine setae (bristles) with fragile bases and a row of barbs on the tip. The barbs cause intense irritation but there is no evidence that they carry any kind of venom. A few defend themselves against wasps by including networks of very robust threads in their webs, giving the spider time to flee while the wasps are struggling with the obstacles. The golden wheeling spider, Carparachne aureoflava, of the Namibian desert escapes parasitic wasps by flipping onto its side and cartwheeling down sand dunes.

 

SOCIAL SPIDERS

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. The genus Anelosimus has a strong tendency towards sociality: all known American species are social, and species in Madagascar are at least somewhat social. Members of other species in the same family but several different genera have independently developed social behavior. For example, although Theridion nigroannulatum belongs to a genus with no other social species, T. nigroannulatum build colonies that may contain several thousand individuals that co-operate in prey capture and share food. Other communal spiders include several Philoponella species (family Uloboridae), Agelena consociata (family Agelenidae) and Mallos gregalis (family Dictynidae). Social predatory spiders need to defend their prey against kleptoparasites ("thieves"), and larger colonies are more successful in this. The herbivorous spider Bagheera kiplingi lives in small colonies which help to protect eggs and spiderlings. Even widow spiders (genus Latrodectus), which are notoriously cannibalistic, have formed small colonies in captivity, sharing webs and feeding together.

 

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 direction required to build them may have been inherited from the common ancestors of most spider groups. However, the majority of spiders build non-orb webs. It used to be thought that the sticky orb web was an evolutionary innovation resulting in the diversification of the Orbiculariae. Now, however, it appears that non-orb spiders are a sub-group 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.

 

ORB WEBS

About half the potential prey that hit orb webs escape. A web has to perform three functions: intercepting the prey (intersection), absorbing its momentum without breaking (stopping), and trapping the prey by entangling it or sticking to it (retention). No single design is best for all prey. For example: wider spacing of lines will increase the web's area and hence its ability to intercept prey, but reduce its stopping power and retention; closer spacing, larger sticky droplets and thicker lines would improve retention, but would make it easier for potential prey to see and avoid the web, at least during the day. However, there are no consistent differences between orb webs built for use during the day and those built for use at night. In fact, there is no simple relationship between orb web design features and the prey they capture, as each orb-weaving species takes a wide range of prey.

 

The hubs of orb webs, where the spiders lurk, are usually above the center, as the spiders can move downwards faster than upwards. If there is an obvious direction in which the spider can retreat to avoid its own predators, the hub is usually offset towards that direction.

 

Horizontal orb webs are fairly common, despite being less effective at intercepting and retaining prey and more vulnerable to damage by rain and falling debris. Various researchers have suggested that horizontal webs offer compensating advantages, such as reduced vulnerability to wind damage; reduced visibility to prey flying upwards, because of the back-lighting 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.

 

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; "ladder-like" webs that appear most effective in catching moths. However, the significance of many variations is unclear.

 

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.

 

TANGLEWEB SPIDERS (COBWEB SPIDERS)

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.

 

OTHER TYPES OF WEBS

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.

 

EVOLUTION

FOSSIL RECORD

Although the fossil record of spiders is considered poor, almost 1000 species have been described from fossils. Because spiders' bodies are quite soft, the vast majority of fossil spiders have been found preserved in amber. 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 prey, producing silk and possibly caring for their young. In a few cases, amber has preserved spiders' egg sacs and webs, occasionally with prey attached; the oldest fossil web found so far is 100 million years old. Earlier spider fossils come from a few lagerstätten, places where conditions were exceptionally suited to preserving fairly soft tissues.

 

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. 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. 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. 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. Its body length is almost 25 mm.

 

Several Carboniferous spiders were members of the Mesothelae, a primitive group now represented only by the Liphistiidae. The mesothelid Paleothele montceauensis, from the Late Carboniferous over 299 million years ago, had five spinnerets. 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.

 

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.

 

WIKIPEDIA

Photographed at the Lexington Wildlife Management Area, Oklahoma, on 7 July 2019.

 

Photographs and text © Bryan Reynolds

All rights reserved. Contact: nature_photo_man@hotmail.com

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Giant house spider

From Wikipedia, the free encyclopedia

Jump to navigationJump to search

Giant house spider

Hausspinne Tegenaria atrica.jpg

Scientific classificationedit

Kingdom:Animalia

Phylum:Arthropoda

Subphylum:Chelicerata

Class:Arachnida

Order:Araneae

Infraorder:Araneomorphae

Family:Agelenidae

Genus:Eratigena

Species:E. atrica

Binomial name

Eratigena atrica

(C. L. Koch, 1843)[1]

Distribution.tegenaria.atrica.1.png

Synonyms[1]

Philoica atrica (C. L. Koch, 1843)

Tegenaria atrica C. L. Koch, 1843

Tegenaria derouetae Denis, 1959

Tegenaria deroueti Dresco, 1957

Tegenaria duellica Simon, 1875

Tegenaria gigantea Chamberlin & Ivie, 1935

Tegenaria hibernica O. Pickard-Cambridge, 1891

Tegenaria larva Simon, 1875

Tegenaria nervosa Simon, 1870

Tegenaria praegrandis Fox, 1937

Tegenaria propinqua Locket, 1975

Tegenaria saeva Blackwall, 1844

The giant house spider, now with the scientific name Eratigena atrica, is one of the biggest spiders of Central and Northern Europe. It was previously placed in the genus Tegenaria, where in addition to Tegenaria atrica, it was also documented as Tegenaria duellica, Tegenaria gigantea and Tegenaria saeva, among others, all thought to be different species. It is now a member of the newly described genus Eratigena.[2] The bite of this species does not pose a threat to humans or pets, and the spider is generally reluctant to bite, preferring instead to hide or escape.

  

female creating egg sac

 

Contents

1Description

2Taxonomy

3Distribution and habitat

4Biology and behavior

5Relationship with Eratigena agrestis

6In popular culture

7References

8Further reading

9External links

Description

 

This section includes a list of references, related reading or external links, but its sources remain unclear because it lacks inline citations. Please help to improve this section by introducing more precise citations. (April 2016) (Learn how and when to remove this template message)

The two sexes do not differ in coloration or markings. Its coloration is mainly dark brown. On its sternum is a lighter marking, with three light spots on each side. The opisthosoma features a lighter middle line with six "spots" on each side. The giant house spider has the same coloration as the domestic house spider, Tegenaria domestica; it has earthy tones of brown and muddy red or yellow. They also have conspicuously hairy legs, palps and abdomen. Despite its English name, this species is not the largest species in the genus (that being Tegenaria parietina). The female body size can reach 18.5 millimetres (0.73 in) in length, with males having a slightly smaller body at around 12 to 15 millimetres (0.47 to 0.59 in) in length. The female leg span is typically around 45 millimetres (1.8 in). The leg span of the male is highly variable, with spans between 25 to 75 millimetres (0.98 to 2.95 in) being common.

 

Its eight eyes are of equal size and are arranged in two rows. As the eyes contain fewer than 400 visual cells, E. atrica can probably only distinguish light and dark.

 

Taxonomy

The first description of a spider now assigned to this species was by Carl Ludwig Koch in 1843, under the name Tegenaria atrica. Other supposedly different species were described later, including Tegenaria saeva by John Blackwall in 1844, Tegenaria duellica by Eugène Simon in 1875 and Tegenaria gigantea by Ralph Vary Chamberlin and Wilton Ivie in 1935. T. gigantea was synonymized with T. duellica in 1978. The three remaining taxa have been regarded as distinct species, particularly in Britain.[2] Thus Roberts (1995) provides distinguishing characters for T. atrica, T. duellica and T. saeva,[3] as does Oxford (2008) for T. duellica (as T. gigantea) and T. saeva.[4] Others consider these three as part of a single morphologically variable species, for which the oldest name, and hence the senior synonym, is T. atrica.[2]

 

A phylogenetic study in 2013 concluded that Tegenaria, as then defined, was not monophyletic, and split off some species, including T. atrica, into the newly created segregate genus Eratigena.[2]

 

Distribution and habitat

E. atrica is found in Europe, Central Asia and Northern Africa. It was unwittingly introduced to the Pacific Northwest of North America circa 1900 due to human activity and has strongly increased in numbers for the last century.[citation needed]

 

In the last few years the spider has been found in several European countries in which it was previously not recorded, like Estonia, Latvia and Lithuania. It is recorded in the checklist of Danish spider species,[5] and is also found in Iceland.[6]

 

The giant house spider's original habitat consists mostly of caves, or dry forests where it is found under rocks, but it is a common spider in people's homes.[citation needed]

 

Biology and behavior

 

Spiderlings

The webs built by the giant house spider are flat and messy with a funnel at one end. They do not contain sticky threads. The spider lurks in the funnel until a small invertebrate happens to get trapped in the web, at which point the spider runs out and attacks it. They usually build their webs in corners (on both the floor and ceiling), between boxes in basements, behind cupboards, in attics, or any other area that is rarely disturbed by large animals or humans. Often found near window openings.[citation needed]

 

E. atrica normally lives for two or three years, but lifetimes of up to six years have been observed. While the female only leaves its nest to feed, males can often be seen wandering around houses during the late summer and early autumn looking for a mate. Males can be found from July to October, adult females occur all year.[citation needed]

 

At least 60 spiderlings emerge from an egg sac. Unusual for spiders, they are subsocial at this stage: they remain together for about a month, but do not cooperate in prey capture. The amount of cannibalism correlates with the amount of available food.[7] E. atrica molts seven or eight times before reaching the immature adult state, and after a final molt reaches maturity.[8]

 

Like most spiders, the spider possesses venom to subdue its prey. Since E. atrica bites can penetrate human skin on occasion, the effects of agatoxin might be felt by bite victims, though these spiders will not bite unless provoked.[citation needed]

 

With speeds clocked at 1.73 ft/s (0.53 m/s; 1.18 mph; 1.90 km/h), the giant house spider held the Guinness Book of World Records for top spider speed until 1987 when it was displaced by solifugids, although the latter are not true spiders.[9]

 

Relationship with Eratigena agrestis

 

E. atrica can attain a leg span of up to 4 inches (100 mm). This specimen is approximately 3 inches (76 mm).

 

A moulting E. atrica

A population of giant house spiders is popularly thought to be a deterrent to the establishment of Eratigena agrestis, known in North America as the "hobo spider", and considered by some to be more likely to bite humans. Giant house spiders may compete with hobo spiders for the same resources. Hobo spiders grow no more than a body size of 15 millimetres (0.59 in) long whereas the larger female giant house spider can have a body size of 18 millimetres (0.71 in),[10] but has proportionately much longer legs.[11]

 

In popular culture

Humorist David Sedaris has written about his relationship with E. atrica. His essay "April In Paris" documents his growing affection towards and domestic association with giant house spiders, particularly one named April.[12] The essay can be found in the collection When You Are Engulfed in Flames.

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Giant house spider

From Wikipedia, the free encyclopedia

Jump to navigationJump to search

Giant house spider

Hausspinne Tegenaria atrica.jpg

Scientific classificationedit

Kingdom:Animalia

Phylum:Arthropoda

Subphylum:Chelicerata

Class:Arachnida

Order:Araneae

Infraorder:Araneomorphae

Family:Agelenidae

Genus:Eratigena

Species:E. atrica

Binomial name

Eratigena atrica

(C. L. Koch, 1843)[1]

Distribution.tegenaria.atrica.1.png

Synonyms[1]

Philoica atrica (C. L. Koch, 1843)

Tegenaria atrica C. L. Koch, 1843

Tegenaria derouetae Denis, 1959

Tegenaria deroueti Dresco, 1957

Tegenaria duellica Simon, 1875

Tegenaria gigantea Chamberlin & Ivie, 1935

Tegenaria hibernica O. Pickard-Cambridge, 1891

Tegenaria larva Simon, 1875

Tegenaria nervosa Simon, 1870

Tegenaria praegrandis Fox, 1937

Tegenaria propinqua Locket, 1975

Tegenaria saeva Blackwall, 1844

The giant house spider, now with the scientific name Eratigena atrica, is one of the biggest spiders of Central and Northern Europe. It was previously placed in the genus Tegenaria, where in addition to Tegenaria atrica, it was also documented as Tegenaria duellica, Tegenaria gigantea and Tegenaria saeva, among others, all thought to be different species. It is now a member of the newly described genus Eratigena.[2] The bite of this species does not pose a threat to humans or pets, and the spider is generally reluctant to bite, preferring instead to hide or escape.

  

female creating egg sac

 

Contents

1Description

2Taxonomy

3Distribution and habitat

4Biology and behavior

5Relationship with Eratigena agrestis

6In popular culture

7References

8Further reading

9External links

Description

 

This section includes a list of references, related reading or external links, but its sources remain unclear because it lacks inline citations. Please help to improve this section by introducing more precise citations. (April 2016) (Learn how and when to remove this template message)

The two sexes do not differ in coloration or markings. Its coloration is mainly dark brown. On its sternum is a lighter marking, with three light spots on each side. The opisthosoma features a lighter middle line with six "spots" on each side. The giant house spider has the same coloration as the domestic house spider, Tegenaria domestica; it has earthy tones of brown and muddy red or yellow. They also have conspicuously hairy legs, palps and abdomen. Despite its English name, this species is not the largest species in the genus (that being Tegenaria parietina). The female body size can reach 18.5 millimetres (0.73 in) in length, with males having a slightly smaller body at around 12 to 15 millimetres (0.47 to 0.59 in) in length. The female leg span is typically around 45 millimetres (1.8 in). The leg span of the male is highly variable, with spans between 25 to 75 millimetres (0.98 to 2.95 in) being common.

 

Its eight eyes are of equal size and are arranged in two rows. As the eyes contain fewer than 400 visual cells, E. atrica can probably only distinguish light and dark.

 

Taxonomy

The first description of a spider now assigned to this species was by Carl Ludwig Koch in 1843, under the name Tegenaria atrica. Other supposedly different species were described later, including Tegenaria saeva by John Blackwall in 1844, Tegenaria duellica by Eugène Simon in 1875 and Tegenaria gigantea by Ralph Vary Chamberlin and Wilton Ivie in 1935. T. gigantea was synonymized with T. duellica in 1978. The three remaining taxa have been regarded as distinct species, particularly in Britain.[2] Thus Roberts (1995) provides distinguishing characters for T. atrica, T. duellica and T. saeva,[3] as does Oxford (2008) for T. duellica (as T. gigantea) and T. saeva.[4] Others consider these three as part of a single morphologically variable species, for which the oldest name, and hence the senior synonym, is T. atrica.[2]

 

A phylogenetic study in 2013 concluded that Tegenaria, as then defined, was not monophyletic, and split off some species, including T. atrica, into the newly created segregate genus Eratigena.[2]

 

Distribution and habitat

E. atrica is found in Europe, Central Asia and Northern Africa. It was unwittingly introduced to the Pacific Northwest of North America circa 1900 due to human activity and has strongly increased in numbers for the last century.[citation needed]

 

In the last few years the spider has been found in several European countries in which it was previously not recorded, like Estonia, Latvia and Lithuania. It is recorded in the checklist of Danish spider species,[5] and is also found in Iceland.[6]

 

The giant house spider's original habitat consists mostly of caves, or dry forests where it is found under rocks, but it is a common spider in people's homes.[citation needed]

 

Biology and behavior

 

Spiderlings

The webs built by the giant house spider are flat and messy with a funnel at one end. They do not contain sticky threads. The spider lurks in the funnel until a small invertebrate happens to get trapped in the web, at which point the spider runs out and attacks it. They usually build their webs in corners (on both the floor and ceiling), between boxes in basements, behind cupboards, in attics, or any other area that is rarely disturbed by large animals or humans. Often found near window openings.[citation needed]

 

E. atrica normally lives for two or three years, but lifetimes of up to six years have been observed. While the female only leaves its nest to feed, males can often be seen wandering around houses during the late summer and early autumn looking for a mate. Males can be found from July to October, adult females occur all year.[citation needed]

 

At least 60 spiderlings emerge from an egg sac. Unusual for spiders, they are subsocial at this stage: they remain together for about a month, but do not cooperate in prey capture. The amount of cannibalism correlates with the amount of available food.[7] E. atrica molts seven or eight times before reaching the immature adult state, and after a final molt reaches maturity.[8]

 

Like most spiders, the spider possesses venom to subdue its prey. Since E. atrica bites can penetrate human skin on occasion, the effects of agatoxin might be felt by bite victims, though these spiders will not bite unless provoked.[citation needed]

 

With speeds clocked at 1.73 ft/s (0.53 m/s; 1.18 mph; 1.90 km/h), the giant house spider held the Guinness Book of World Records for top spider speed until 1987 when it was displaced by solifugids, although the latter are not true spiders.[9]

 

Relationship with Eratigena agrestis

 

E. atrica can attain a leg span of up to 4 inches (100 mm). This specimen is approximately 3 inches (76 mm).

 

A moulting E. atrica

A population of giant house spiders is popularly thought to be a deterrent to the establishment of Eratigena agrestis, known in North America as the "hobo spider", and considered by some to be more likely to bite humans. Giant house spiders may compete with hobo spiders for the same resources. Hobo spiders grow no more than a body size of 15 millimetres (0.59 in) long whereas the larger female giant house spider can have a body size of 18 millimetres (0.71 in),[10] but has proportionately much longer legs.[11]

 

In popular culture

Humorist David Sedaris has written about his relationship with E. atrica. His essay "April In Paris" documents his growing affection towards and domestic association with giant house spiders, particularly one named April.[12] The essay can be found in the collection When You Are Engulfed in Flames.

A night visitor to our bathroom 26/9/2019 TL005287

Giant house spider

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Giant house spider

Hausspinne Tegenaria atrica.jpg

Scientific classificationedit

Kingdom:Animalia

Phylum:Arthropoda

Subphylum:Chelicerata

Class:Arachnida

Order:Araneae

Infraorder:Araneomorphae

Family:Agelenidae

Genus:Eratigena

Species:E. atrica

Binomial name

Eratigena atrica

(C. L. Koch, 1843)[1]

Distribution.tegenaria.atrica.1.png

Synonyms[1]

Philoica atrica (C. L. Koch, 1843)

Tegenaria atrica C. L. Koch, 1843

Tegenaria derouetae Denis, 1959

Tegenaria deroueti Dresco, 1957

Tegenaria duellica Simon, 1875

Tegenaria gigantea Chamberlin & Ivie, 1935

Tegenaria hibernica O. Pickard-Cambridge, 1891

Tegenaria larva Simon, 1875

Tegenaria nervosa Simon, 1870

Tegenaria praegrandis Fox, 1937

Tegenaria propinqua Locket, 1975

Tegenaria saeva Blackwall, 1844

The giant house spider, now with the scientific name Eratigena atrica, is one of the biggest spiders of Central and Northern Europe. It was previously placed in the genus Tegenaria, where in addition to Tegenaria atrica, it was also documented as Tegenaria duellica, Tegenaria gigantea and Tegenaria saeva, among others, all thought to be different species. It is now a member of the newly described genus Eratigena.[2] The bite of this species does not pose a threat to humans or pets, and the spider is generally reluctant to bite, preferring instead to hide or escape.

  

female creating egg sac

 

Contents

1Description

2Taxonomy

3Distribution and habitat

4Biology and behavior

5Relationship with Eratigena agrestis

6In popular culture

7References

8Further reading

9External links

Description

 

This section includes a list of references, related reading or external links, but its sources remain unclear because it lacks inline citations. Please help to improve this section by introducing more precise citations. (April 2016) (Learn how and when to remove this template message)

The two sexes do not differ in coloration or markings. Its coloration is mainly dark brown. On its sternum is a lighter marking, with three light spots on each side. The opisthosoma features a lighter middle line with six "spots" on each side. The giant house spider has the same coloration as the domestic house spider, Tegenaria domestica; it has earthy tones of brown and muddy red or yellow. They also have conspicuously hairy legs, palps and abdomen. Despite its English name, this species is not the largest species in the genus (that being Tegenaria parietina). The female body size can reach 18.5 millimetres (0.73 in) in length, with males having a slightly smaller body at around 12 to 15 millimetres (0.47 to 0.59 in) in length. The female leg span is typically around 45 millimetres (1.8 in). The leg span of the male is highly variable, with spans between 25 to 75 millimetres (0.98 to 2.95 in) being common.

 

Its eight eyes are of equal size and are arranged in two rows. As the eyes contain fewer than 400 visual cells, E. atrica can probably only distinguish light and dark.

 

Taxonomy

The first description of a spider now assigned to this species was by Carl Ludwig Koch in 1843, under the name Tegenaria atrica. Other supposedly different species were described later, including Tegenaria saeva by John Blackwall in 1844, Tegenaria duellica by Eugène Simon in 1875 and Tegenaria gigantea by Ralph Vary Chamberlin and Wilton Ivie in 1935. T. gigantea was synonymized with T. duellica in 1978. The three remaining taxa have been regarded as distinct species, particularly in Britain.[2] Thus Roberts (1995) provides distinguishing characters for T. atrica, T. duellica and T. saeva,[3] as does Oxford (2008) for T. duellica (as T. gigantea) and T. saeva.[4] Others consider these three as part of a single morphologically variable species, for which the oldest name, and hence the senior synonym, is T. atrica.[2]

 

A phylogenetic study in 2013 concluded that Tegenaria, as then defined, was not monophyletic, and split off some species, including T. atrica, into the newly created segregate genus Eratigena.[2]

 

Distribution and habitat

E. atrica is found in Europe, Central Asia and Northern Africa. It was unwittingly introduced to the Pacific Northwest of North America circa 1900 due to human activity and has strongly increased in numbers for the last century.[citation needed]

 

In the last few years the spider has been found in several European countries in which it was previously not recorded, like Estonia, Latvia and Lithuania. It is recorded in the checklist of Danish spider species,[5] and is also found in Iceland.[6]

 

The giant house spider's original habitat consists mostly of caves, or dry forests where it is found under rocks, but it is a common spider in people's homes.[citation needed]

 

Biology and behavior

 

Spiderlings

The webs built by the giant house spider are flat and messy with a funnel at one end. They do not contain sticky threads. The spider lurks in the funnel until a small invertebrate happens to get trapped in the web, at which point the spider runs out and attacks it. They usually build their webs in corners (on both the floor and ceiling), between boxes in basements, behind cupboards, in attics, or any other area that is rarely disturbed by large animals or humans. Often found near window openings.[citation needed]

 

E. atrica normally lives for two or three years, but lifetimes of up to six years have been observed. While the female only leaves its nest to feed, males can often be seen wandering around houses during the late summer and early autumn looking for a mate. Males can be found from July to October, adult females occur all year.[citation needed]

 

At least 60 spiderlings emerge from an egg sac. Unusual for spiders, they are subsocial at this stage: they remain together for about a month, but do not cooperate in prey capture. The amount of cannibalism correlates with the amount of available food.[7] E. atrica molts seven or eight times before reaching the immature adult state, and after a final molt reaches maturity.[8]

 

Like most spiders, the spider possesses venom to subdue its prey. Since E. atrica bites can penetrate human skin on occasion, the effects of agatoxin might be felt by bite victims, though these spiders will not bite unless provoked.[citation needed]

 

With speeds clocked at 1.73 ft/s (0.53 m/s; 1.18 mph; 1.90 km/h), the giant house spider held the Guinness Book of World Records for top spider speed until 1987 when it was displaced by solifugids, although the latter are not true spiders.[9]

 

Relationship with Eratigena agrestis

 

E. atrica can attain a leg span of up to 4 inches (100 mm). This specimen is approximately 3 inches (76 mm).

 

A moulting E. atrica

A population of giant house spiders is popularly thought to be a deterrent to the establishment of Eratigena agrestis, known in North America as the "hobo spider", and considered by some to be more likely to bite humans. Giant house spiders may compete with hobo spiders for the same resources. Hobo spiders grow no more than a body size of 15 millimetres (0.59 in) long whereas the larger female giant house spider can have a body size of 18 millimetres (0.71 in),[10] but has proportionately much longer legs.[11]

 

In popular culture

Humorist David Sedaris has written about his relationship with E. atrica. His essay "April In Paris" documents his growing affection towards and domestic association with giant house spiders, particularly one named April.[12] The essay can be found in the collection When You Are Engulfed in Flames.

852

Seen in my porch front leg to back leg 3 inches and a 1 inch body.

The spider species Tegenaria domestica, commonly known as the barn funnel weaver in North America and the domestic house spider in Europe, is a member of the funnel-web family Agelenidae

Like all members of the funnel weaver family Agelenidae, these spiders spin dense, non-sticky, sheet-like webs with a funnel-like retreat where the spider hides.

 

Male, 15mm length

Family Agelenidae.

Sevastopol, Crimea.

 

Паук-воронкопряд, ♀.

Female funnel weaver, family Agelenidae, at night. Leavenworth, Kansas, USA, August 20, 2017.

North American Hobo Spider (Agelenidae) with Large, Flat Funnel Web, Summer Morning, Rocky Mountain Front Range, Colorado

 

Phylum Arthropoda

Class Arachnida

Family Agelenidae

Genus Eratigena

Species E. agrestis

“Grass Spiders” are represented by 13 species collectively found throughout most of the U.S. and southern Canada, and northern Mexico. Like all members of the funnel weaver family Agelenidae, they spin dense, non-sticky, sheet-like webs with a funnel-like retreat where the spider hides.

This spider has a nice inviting bejeweled parlor just waiting for an unsuspecting guest.

 

Funnel-web spider. There are more than 1146 species. They have a special spatial perception that allows them to orient themselves and navigate even in complete darkness. wikipedia

 

Funnel-web spider (Family Agelenidae)

North American Hobo Spider (Agelenidae) with Large, Flat Funnel Web, Summer Morning, Rocky Mountain Front Range, Colorado

 

Phylum Arthropoda

Class Arachnida

Family Agelenidae

Genus Eratigena

Species E. agrestis

Female on large sheet web amongst heather from which this animal gets its name, in the family Agelenidae

Mature male funnel weaver, family Agelenidae. Leavenworth, Kansas, USA, August 23, 2017.

Agelenopsis spp

Summary

“Grass Spiders” are represented by 13 species collectively found throughout most of the U.S. and southern Canada, and northern Mexico. Like all members of the funnel weaver family Agelenidae, they spin dense, non-sticky, sheet-like webs with a funnel-like retreat where the spider hides.

 

Phylum: Arthropoda LATREILLE, 1829 (arthropods, Gliederfüßer)

Subphylum: Chelicerata

Class: Arachnida CUVIER, 1812 (arachnids, Spinnentiere)

Subclass: Micrura

Infraclass: Megoperculata

Order: Araneae CLERCK, 1757 (spiders)

Suborder: Araneomorphae SIMON, 1892 (Echte Webspinnen)

Superfamily: Agelenoidea

Family: Agelenidae C. L. KOCH, 1837 (funnelweb spiders, Trichterspinnen)

Genus: Eratigena BOLZERN, BURCKHARDT & HÄNGGI, 2013 [Syn. Tegenaria]

Eratigena atrica C. L. KOCH, 1843 (Giant House Spider)

 

Central Germany, N-Hesse, Kassel: Dönche (NP), 225m asl., 18.08.2016

 

IMG_9364

The Giant House Spider is common in and around buildings.. The webs can be found in dark corners of rooms, garages, sheds, under rocks and logs, etc. This species is often found inside bathtubs and sinks, having gotten stuck while trying to climb in for a drink of water.

Constructs a web characteristic of most members of the family Agelenidae: a funnel-shaped retreat with a radiating flat sheet of webbing. Spider typically remains inside the retreat during the day, motionless, unless prey enters the web; at which point, it will then rush out to grab it. At night, the spider may stay perched out on the main sheet of webbing to await prey. It will dart back into the retreat if disturbed.

Agelenopsis sp. (Wikipedia)

(Encyclopedia Of Life) (Bug Guide)

Anybody on my doorstep i can eat ?

(Greenville Cypress Preserve)

(Greenville) (Mississippi)

Might be a Agelenopsis naevia...

But we are not 100 % sure !

Savage/Christmas Creek Preserve, Orange County, FL, January 2020.

Funnel-web weavers like this one (family Agelenidae) frequently build their webs high up in shrubs, trees, and on buildings. This afford the occasional ventral veiw. Cheyenne Mountain State Park, Colorado, USA, August 11, 2016.

This lady was sooo fast catching her fly...

She must have been pretty hungry !!

Agelena labyrinthica ♀ (Wikipedia)

(Encyclopedia Of Life) (Eurospiders)

Spiders that build funnel-shaped webs are usually known as funnel-web spiders, or agelenidae. There are more than 80 species of funnel-webs. These spiders are commonly found in North America and are mostly harmless to humans. There are, however, spiders in Australia that also produce funnel-shaped webs and can be very dangerous to humans. These spiders actually belong to another family, called hexathelidae, and are not related to North American funnel-webs. eHow.com

Down on Sherkin Island during the summer and rushing to catch up with the youngsters I spotted this spider. My first thought was that it was a "Funnel Web Spider". It is not very sharp, indeed it is not sharp at all, but it does give some idea of the little creature. He is there sitting at the entrance to his very complex web waiting patiently for some callers. The deeper c.olour around him is a tunnel approx. 2 inches long and surrounded by looser filaments. I looked it up on the web and it is not a FWS as I thought but appears to be a "Labyrinth Spider" It was a lively wee fellow and darted up and down its tunnel at the slightest movement.

Wikipedia has the following:

Agelena labyrinthica is a species of spiders in the family Agelenidae. It is a widespread species in Europe.

A. labyrinthica build flat plate surface webs connected to funnel-shaped retreats similar to labyrinths, which are typically constructed between low lying grass and vegetation.[1] These webs can be at ground level, or up to 1.5 metres (4 ft 11 in) from the ground, however, the majority are found approximately 60 centimetres (24 in) off of the ground.[1][3] These spiders are fairly common in Europe and Central Europe, and are typically concentrated in areas near forests and low lying vegetation, as well as in dry grasslands.

Arachtober 22nd, Funnel Weaver Spider, Family Agelenidae - Funnel Weavers, Genus Agelenopsis - Grass Spiders - eating a gob of goo.

For this genus of spiders, the web is a horizontal, sheet-like web, with a small funnel-like tube off to a side and a 3-dimensional barrier web over the top. When a flying insect hits the barrier, it falls into the sheet below. The funnel (or tube) is what the family (Agelenidae) is named for, and is used by the spider for hunting. The spider will lie in wait in the funnel, and when an insect flies hits the barrier and falls on the web, the spider will rush out, very quickly check to see if it is prey, and if it is prey, bite it. The venom is fast-acting on the prey, so once the prey is subdued the spider will drag the prey back into the funnel (for safety while eating, and to prevent other insects from recognizing the danger that lurks below on the web).

 

For Agelenopsis spp. spiders, the web is not sticky. If the insect lands/falls on the web, the web will actually become tangled around the prey's feet, temporarily ensnaring it in the web. The funnel web for Agelenopsis is a distinctive web, and often is noticed in bushes and grass, especially in the early fall mornings, where the dew has collected on the web. The webs can be expansive, covering several square feet, or just small webs in the grass.

 

Raynox DCR-250 mounted on my Panasonic FZ8.

Agelena labyrinthica ♀ (Wikipedia) (Encyclopedia Of Life) (Eurospiders)

Although she's got prey, she keeps an eye (or more) on the lens in front of her..

Night Photography, the Northern Sonoran Desert - 2015.

Native spiders are highly adaptable and opportunistic, this spider built her web in between hedgehog cactus.

Night Photography, the Northern Sonoran Desert - 2015.

Native spiders are highly adaptable and opportunistic, this spider built her web in between hedgehog cactus.

Morning dew on a spider web in the garden. The green is a evergreen needle I broke off the Mugo Pine that the web was woven on. I dropped it next to the 'funnel' where the spider was hiding, hoping to draw the spider out for a photo op. She didn't fall for it.

 

Funnel web weavers (Family Agelenidae) are small to medium sized spiders often found in grassy fields, low shrubbery, or living among leaf litter in forests. They spin sheet webs of nonsticky silk with a characteristic funnel extending off to one side. The funnel is where the spider hides while awaiting prey. There is a 3-dimensional barrier web spun above the sheet web, and when a prey item falls through onto the sheet web, the spider quickly runs out and bites its victim, then drags it back to the funnel to feed. These sheet webs are nearly invisible unless covered with dewdrops on a cool morning, and the spider can move very quickly over the surface. It almost looks as if the spider is walking on air. There are over 400 North American species.

Spiders in the most common genus, Agelenopsis, are commonly called "grass spiders," after their habit of building their combination sheet-and-funnel webs in grass and low shrubs.

 

An old saying about grass spiders: when there is dew on thier webs in the lawn in the morning, it will be a beautiful day.

 

For more incredible photos and information on spiders, including the Funnel Web Weaver spider I was attempting to get an image of, go to: www.cirrusimage.com/spider.htm</a

        

So we found this a few weeks ago running around our basement apartment.

The spider 3''-4'' in diameter (leg spread).

I of course caught it in a large jar and photographed it.

I did some research tonight and learned on close inspection that it is not a hobo spider (as I first thought).

Hobo spiders (Tegenaria agrestis) are venomous (though rarely fatal). Giant house spiders are not, (at least to humans, see comment below) and apparently are thought to out compete the hobo.

 

See the description of the next pics for more info.

 

For the taxonomy nuts:

Kingdom Animalia

Phylum Arthropoda

Subphylum Chelicerata

Class Arachnida

Order Araneae

Suborder Opisthothelae

Infraorder Araneomorphae (True Spiders)

No Taxon (Entelegynes)

Family Agelenidae (Funnel-Web Spiders)

Genus Tegenaria

Species duellica (gigantea to some)

Savage Christmas Creek Preserve, Orange County, FL, January 2013.

CLASS: Chelicerata (Arachnids)

ORDER: Araneae

FAMILY: Agelenidae (Funnel-web Spider)

GENUS: Agelenopsis (Grass Spiders)

SPECIES: **

In my garden. Eating a woodlouse. Zaragoza, Spain

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