Spider webs
Spiders that use silk to capture prey utilize various techniques. Ground-dwelling trap-door spiders construct silk-lined tubes, sometimes with silk trapdoors, from which they dart out to capture passing insects. Other tube-dwelling spiders place silk trip threads around the mouth of the tube. When an insect touches these threads, vibrations inform the spider of a victim’s presence. Funnel-weaving spiders live in silk tubes with a narrow end that extends into vegetation or a crevice and an expanded sheetlike end that vibrates when an insect walks across it. Many web spiders construct silk sheets in vegetation, sometimes one above the other, and often add anchor threads, which trip unsuspecting insects. The irregular three-dimensional web of cobweb spiders (family Theridiidae) has anchoring threads of sticky silk. An insect caught in the web or touching an anchor line becomes entangled, increasingly so if it struggles. If a thread breaks, the elasticity of it pulls the insect toward the centre of the web.
The most elaborate webs are those of the orb weavers, whose circular nets are conspicuous on dewy mornings. This type of web is constructed by several spider families, which suggests that it is an efficient trap that enables the largest area to be covered with the least possible silk. The web acts like an air filter, trapping weak-flying insects that cannot see the fine silk. Most orb webs are rebuilt every day. The web may be up only during the day or only at night. If a web is damaged during capture of prey, the spider will repair that area. The ways by which spiders keep from becoming entangled in their own webs are not completely understood, nor is their mechanism for cutting the extremely elastic silk threads that are used in web construction.
To begin orb-web construction, the spider releases a silk thread that is carried by wind. If the free end does not become attached to an object, the spider may pull it back and feed on it. If it becomes firmly attached—for example, to a twig—the spider secures the thread and crosses the newly formed bridge, reinforcing it with additional threads. The spider then descends from the centre of the bridge, securing a thread on the ground or on a twig. The centre, or hub, of the web is established when the spider returns to the bridge with a thread and carries it partway across the bridge before securing it; this thread is the first radius, or spoke. After all the spokes are in place, the spider returns to the hub and constructs a few temporary spirals of dry silk toward the outside of the web. The spider then reverses direction, deposits ensnaring silk, and removes the initial spiral. The ensnaring threads form a dense spiral. It takes only about an hour to weave the radii and orb.
Some species attach a signal thread from the hub to a retreat in a leaf so that they are informed (by vibrations) of trapped insects; others remain head-down in the centre of the orb, locating prey by sensing tensions or vibrations in individual spokes. Webs of two spider families (Araneidae and Tetragnathidae) have spirals constructed of a sticky material that dries out after several days and must be rebuilt.
Spiders of the family Uloboridae build a web of woolly (cribellate) ensnaring silk. One group within this family (genus Hyptiotes) weaves only a partial orb. The spider, attached by a thread to vegetation, holds one thread from the tip of the hub until an insect brushes the web. The spider then alternately relaxes and tightens the thread, and the struggling victim becomes completely entangled. Tiny theridiosomatid spiders also control web tension.
Ogre-faced spiders (family Deinopidae) build small flat webs during the evening hours and then cut the attachments and spread the web among their four long front legs. During the night the web is thrown over a passing insect. The spider abandons or eats the web in the morning and passes the day resting on a branch before constructing a new web.
Bolas spiders (Mastophora, Ordgarius) release a single thread with a sticky droplet at the end and hold it with one leg. Some species swing this “bola,” and others throw it when a moth approaches. Male moths are attracted to this spider by its odour, which mimics that of female moths. Many other examples of web specializations have been described.
Spiders usually wrap a captured insect in silk while turning it, as on a spit, before biting it and carrying it either to a retreat or to the hub of the web for feeding or storage. Although the detachable scales of butterfly and moth wings facilitate their escape from the web, spiders have evolved a counterstrategy: they bite before wrapping them rather than afterward.
Some tropical species of spiders are social and live in large communal webs containing hundreds of individuals, most of them female. They cooperate to build and repair the web. The pack of spiders subdues, kills, and consumes insects that have been caught in the communal web.
Classification
Distinguishing taxonomic features
The Araneida are separated into three suborders: Mesothelae (segmented spiders), Orthognatha (mygalomorph spiders), and Labidognatha (araneomorph spiders). The segmented spiders are easily distinguished by indentations on the top of the abdomen—evidence of spiders’ common ancestry with scorpions. The other two suborders are differentiated on the basis of the type of movement of the two jaws; i.e., movement forward and down is orthognath (paraxial), and movement sideways and together is labidognath (diaxial). Other external features that distinguish suborders include the structure of the male pedipalps and the presence or absence of an epigynum in the female. Internal differentiating features include the presence and number of book lungs, number of small openings (ostia) in the heart, and extent of fusion of nerve ganglia in the prosoma. Families are distinguished on the basis of such characteristics as number and spacing of simple eyes, number of tarsal claws, number of spinnerets, habits, structure of chelicerae, and specialized (apomorph) characters such as glands, setae, and teeth and peculiarities of the external genitalia. Species and also genera of araneomorph spiders are usually separated by specializations of the female epigynum and male pedipalp.
Annotated classification
Numerous classification schemes were published in the 1930s, most of them in response to one by Alexander Petrunkevitch, but none of these is now acceptable and up-to-date. All classifications have relied heavily on the work of Eugène Simon, who published in France in the late 19th century. Newer tools, such as scanning electron microscopy, molecular methods, and cladistics, remain little-tried for spiders, but they have changed traditional classification schemes. In addition, many new spiders have been found in the Southern Hemisphere that do not readily fit into established families, a situation that prompts the proposal of new ones, though without an overall view for a new system. Fewer than 30 percent of the large neotropical spiders are known (and probably fewer of the small neotropical spiders), while 80 percent or more of the species in northern and central Europe, northern North America, Korea, and Japan are known. The known species are placed into about 110 families, only the most important of which are described below.
- Order Araneida or Araneae (spiders)
- More than 46,700 species described in about 110 families virtually worldwide. Arachnids with an unsegmented abdomen (opisthosoma) attached to cephalothorax (prosoma) by pedicel; prosoma with poison glands whose ducts open through large chelicerae with a movable fang, no pincers; leglike pedipalps usually with modifications (endites) on coxae to aid in feeding, modified as copulatory organs in males; abdomen usually with 2 pairs of respiratory organs, anterior book lungs and posterior tracheae; spinnerets at posterior end of abdomen; 4 pairs of walking legs; 1 or 2 pairs of coxal glands.
- Suborder Labidognatha (araneomorph spiders)
- Chelicerae labidognath (diaxial), attached below carapace; pedipalpal coxae with endites; usually 1 pair of book lungs, sometimes replaced by tracheae; heart with 3, sometimes 2, ostia; 13th through 18th ganglia lost, others fused.
- Family Salticidae (jumping spiders)
- More than 5,800 species; mostly tropical, but many species also found in northern and Arctic regions. Hunt during daylight hours by stalking and jumping on prey; best vision of all spiders.
- Family Linyphiidae (sheet-web weavers and dwarf spiders)
- 4,300 species worldwide. Chelicerae with many teeth, often with stridulating (sound-producing) file on side; legs with many strong setae.
- Family Araneidae (orb weavers)
- 2,800 species worldwide; Araneus widespread. Legs with strong setae; chelicerae strong, truncated proximally, with 6 to 8 teeth; usually construct orb webs; Mastophora, the bolas spider, probably venomous to humans; many species make no webs but instead wait for prey to stumble into them.
- Family Lycosidae (wolf spiders)
- 2,300 species worldwide, including numerous species in the Arctic and on high mountains and the original tarantula, Lycosa tarentula of southern Europe. Eyes in 3 rows, anterior row with 4 eyes; hunting spiders; females carry egg sac attached to spinnerets and carry young on abdomen; may dig tubes in soil.
- Family Theridiidae (cobweb weavers or comb-footed spiders)
- More than 2,200 species worldwide. Comb on each rear leg, used to throw silk over prey (attack wrap); chelicerae pointed on near end under carapace, few teeth; no stridulating structures on side of chelicerae; web irregular; sometimes hunt on ground; Latrodectus (widows) poisonous to mammals.
- Family Thomisidae (crab spiders)
- 2,000 species; common and found worldwide. Often sit on flowers awaiting insects; some change colour; some live on or under bark.
- Family Gnaphosidae
- More than 2,000 common and widespread species. Anterior (lateral) spinnerets cylindrical and separated; posterior median eyes often oval and diagonal; nocturnal hunters.
- Family Sparassidae or Heteropodidae (huntsman spiders, tarantulas in Australia)
- Found in most tropical regions. Eyes in 2 rows; legs extended sideways; large, slightly flattened body.
- Family Tetragnathidae (long-jawed orb weavers)
- 1,000 species worldwide. Males with long chelicerae; epigynum often secondarily lost.
- Family Pholcidae (daddy longlegs spiders)
- About 960 species worldwide. Similar to the nonspiders called daddy longlegs of the order Opiliones. Tarsi of legs with many false articulations; no tracheae; web loose and tangled; Pholcus of Europe and America.
- Family Amaurobiidae
- 680 species common worldwide. Cribellum; 3 tarsal claws without brush of setae; tarsi with dorsal row of trichobothria; resemble Agelenidae; make an irregular funnel web between stones.
- Family Dictynidae
- About 560 species common in temperate areas. Cribellum; 3 tarsal claws; tarsi lack trichobothria and brush of setae; small in size; make irregular webs under leaves or in branches of herbs.
- Family Clubionidae (sac spiders)
- More than 530 species; common and widespread. Hunting spiders; anterior pair of spinnerets conical and touching; Cheiracanthium species somewhat venomous to humans.
- Family Dysderidae
- 500 species worldwide. Respiratory tracheae with 4 spiracles (openings) in 2 pairs, 1 behind the other.
- Family Agelenidae (funnel weavers)
- 500 species worldwide. Eyes in 2 rows; anterior (lateral) spinnerets long; most make a flat funnel web in vegetation and a tube-shaped retreat at the side; Argyroneta, the Eurasian water spider, in its own family but related to agelenids, constructs an air bell beneath the surface of slow-moving waters.
- Family Ctenidae (wandering spiders)
- 440 mainly large tropical species. Eyes in 3 rows; median eyes face same direction, form a trapezoid wider behind than in front; eyes of posterior row largest, anterior laterals smallest; 1st 2 legs armed with strong ventral setae; some adults with only 2 tarsal claws; aggressive Phoneutria fera of South America venomous to humans.
- Family Oxyopidae (lynx spiders)
- 420 species worldwide. Eyes arranged in a hexagon; hunt on vegetation, pounce on prey.
- Family Uloboridae
- About 260 species worldwide. Cribellum; lack poison glands; 3 tarsal claws; eyes in 3 rows; anal tubercle large; make orb webs; Hyptiotes are called triangle spiders.
- Family Scytodidae (spitting spiders)
- 160 species mostly in tropical and subtropical regions. 6 eyes arranged in 3 groups; high, domed carapace, slopes anteriorly; catch prey with well-aimed squirt of saliva.
- Family Sicariidae (six-eyed crab spiders)
- About 130 species of Southern Hemisphere deserts; includes genus Loxosceles (recluse spiders). Large, 6 eyes, low carapace; legs extended toward sides; burrow in sand.
- Family Theridiosomatidae (ray spiders)
- More than 100 species. Globular abdomen; high clypeus (area below eyes); orb webs.
- Family Deinopidae (ogre-faced spiders)
- 60 tropical species. Cribellum; 3 tarsal claws; eyes in 3 rows; anal tubercle large; Dinopis with 2 huge eyes, holds web, throws it over prey.
- Suborder Orthognatha (mygalomorph spiders)
- Most species large and long-lived in warm climates. 2 pairs of book lungs; heart with 4, rarely 3, ostia; bulb of male pedipalps simple; female without epigynum; 13th through 18th ganglia lost, others fused.
- Family Theraphosidae (hairy mygalomorphs, tarantulas, baboon spiders)
- 900 mostly tropical species including tarantulas of North America, baboon spiders of South Africa, and bird-eating spiders of Australia, New Guinea, and South America. 2 tarsal claws; large, hairy, nonsocial, nocturnal; some burrow, others inhabit trees.
- Family Dipluridae (diplurid funnel-web mygalomorphs)
- 188 mostly tropical species. 3 tarsal claws; 4 to 6 spinnerets, posterior (lateral) pair very long; body hairy; web similar to funnel webs of Agelenidae.
- Family Ctenizidae (ctenizid trap-door spiders)
- 128 mostly tropical species. Chelicerae with structure (rake or rastellum) used to dig; 3 tarsal claws; eyes closely grouped; most species at least 3 cm or more in length; inhabit silk-lined tubes in ground, with entrances covered by hinged silk lids.
- Family Hexathelidae
- 85 mostly tropical species. Arched, glabrous carapace differentiates it from Dipluridae; funnel-web spiders (genus Atrax) of southeastern Australia are venomous.
- Family Atypidae (purse-web spiders)
- 43 species of Europe, North America, Japan, Myanmar, and Java. 3 tarsal claws; 6 spinnerets; less than 3 cm long; live in closed silk tubes partly below ground; bite prey through tube and pull it in.
- Suborder Mesothelae (segmented spiders)
- About 100 species in 1 family, Liphistiidae, found from Japan to Southeast Asia. Inhabit trapdoor tubes in ground; remnants of abdominal segmentation clearly visible dorsally from 7th segment (pedicel) to 18th; 8 spinnerets at middle of abdomen; male pedipalps relatively complicated; epigynum absent in females; 18 ganglia in prosoma more distinctly separated than those in other suborders; heart with 5 pairs of ostia in segments 8 to 13; 2 pairs of book lungs.