Spider (order Araneae)

 Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs
generally able to inject venom, and spinnerets that extrude silk. They are the largest order of
arachnids and rank seventh in total species diversity among all orders of organisms. Spiders are
found worldwide on every continent except for Antarctica, and have become established in nearly
every habitat with the exceptions of air and sea colonization. As of August 2021, 49,623 spider
species in 129 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 (as
with all arachnids) differ from other arthropods in that the usual body segments are fused into two
tagmata, the prosoma, or cephalothorax, and opisthosoma, or abdomen, and joined by a small,
cylindrical pedicel (however, as there is currently neither paleontological nor embryological
evidence that spiders ever had a separate thorax-like division, there exists an argument against the
validity of the term cephalothorax, which means fused cephalon (head) and the thorax. Similarly,
arguments can be formed against use of the term abdomen, as the opisthosoma of all spiders
contains a heart and respiratory organs, organs atypical of an abdomen). Unlike 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.
 Body plan
 Spiders are chelicerates and therefore arthropods. As arthropods they have: segmented bodies with jointed
limbs, all covered in a cuticle made of chitin and proteins; heads that are composed of several segments that
fuse during the development of the embryo. Being chelicerates, their bodies consist of two tagmata, sets of
segments that serve similar functions: the foremost one, called the cephalothorax or prosoma, is a complete
fusion of the segments that in an insect 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 abdome
 Eyes
 Spiders have primarily four pairs of eyes on the top-front area of the cephalothorax,
arranged in patterns that vary from one family to another. The principal pair at the
front are of the type called pigment-cup ocelli ("little eyes"), which in most arthropods
are only capable of detecting the direction from which light is coming, using the
shadow cast by the walls of the cup. However, in spiders these eyes are capable of
forming images. The other pairs, called secondary eyes, are thought to be derived from
the compound eyes of the ancestral chelicerates, but no longer have the separate
facets typical of compound eyes. Unlike the principal eyes, in many spiders these
secondary eyes detect light reflected from a reflective tapetum lucidum, and wolf
spiders can be spotted by torchlight reflected from the tapeta. On the other hand, the
secondary eyes of jumping spiders have no tapeta. Other differences between the
principal and secondary eyes are that the latter have rhabdomeres that point away
from incoming light, just like in vertebrates, while the arrangement is the opposite in
the former. The principal eyes are also the only ones with eye muscles, allowing them
to move the retina. Having no muscles, the secondary eyes are immobile. The visual
acuity of some jumping spiders exceeds by a factor of ten that of dragonflies, which
have by far the best vision among insects.[citation needed] This acuity is achieved by a
telephotographic series of lenses, a four-layer retina, and the ability to swivel the eyes
and integrate images from different stages in the scan.[citation needed] The downside
is that the scanning and integrating processes are relatively slow. There are spiders
with a reduced number of eyes, the most common having six eyes (example, Periegops
suterii) with a pair of eyes absent on the anterior median line. Other species have four
eyes and members of the Caponiidae family can have as few as two. Cave dwelling
species have no eyes, or possess vestigial eyes incapable of sight.
 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.[30]
The only extensor muscles in spider legs are located in the three hip joints (bordering
the coxa and the trochanter).[31] As a result, a spider with a punctured cephalothorax
cannot extend its legs, and the legs of dead spiders curl up.[12] Spiders can generate
pressures up to eight times their resting level to extend their legs,[32] 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.[12] 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.[31] Most spiders that hunt
actively, rather than relying on webs, have dense tufts of fine bristles between the
paired claws at the tips of their legs. These tufts, known as scopulae, consist of bristles
whose ends are split into as many as 1,000 branches, and enable spiders with scopulae
to walk up vertical glass and upside down on ceilings. It appears that scopulae get their
grip from contact with extremely thin layers of water on surfaces.[12] Spiders, like
most other arachnids, keep at least four legs on the surface while walking or running.
 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 onto which they ejaculate and then transfer the sperm to special
syringe-styled structures, palpal bulbs or palpal organs, borne on the tips of the 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
via one or two openings on the underside of her abdomen. Female spiders' reproductive tracts are
arranged in one of two ways. The ancestral arrangement ("haplogyne" or "non-entelegyne") consists
of a single genital opening, leading to two seminal receptacles (spermathecae) in which females
store sperm. In the more advanced arrangement ("entelegyne" ), there are two further openings
leading directly to the spermathecae, creating a "flow through" system rather than a "first-in first-
out" one. Eggs are as a general rule only fertilized during oviposition when the stored sperm is
released from its chamber, rather than in the ovarian cavity. A few exceptions exist, such as
Parasteatoda tepidariorum. In these species the female appears to be able to activate the dormant
sperm before oviposition, allowing them to migrate to the ovarian cavity where fertilization occurs.
The only known example of direct fertilization between male and female is an Israeli spider,
Harpactea sadistica, which has evolved traumatic insemination. In this species the male will
penetrate its pedipalps through the female's body wall and inject his sperm directly into her
ovaries, where the embryos inside the fertilized eggs will start to develop before being laid.
 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 subgroup that evolved from orb-web spiders, and non-orb spiders have over 40%
more species and are four times as abundant as orb-web spiders. Their greater success
may be because sphecid wasps, which are often the dominant predators of spiders,
much prefer to attack spiders that have flat webs.
 Orb
About half the potential prey that hit orb webs escape. A web has to perform three functions: intercepting the
prey (intersection), absorbing its momentum without breaking (stopping), and trapping the prey by entangling it
or sticking to it (retention). No single design is best for all prey. For example: wider spacing of lines will increase
the web's area and hence its ability to intercept prey, but reduce its stopping power and retention; closer
spacing, larger sticky droplets and thicker lines would improve retention, but would make it easier for potential
prey to see and avoid the web, at least during the day. However, there are no consistent differences between
orb webs built for use during the day and those built for use at night. In fact, there is no simple relationship
between orb web design features and the prey they capture, as each orb-weaving species takes a wide range of
prey. The hubs of orb webs, where the spiders lurk, are usually above the center, as the spiders can move
downwards faster than upwards. If there is an obvious direction in which the spider can retreat to avoid its own
predators, the hub is usually offset towards that direction. Horizontal orb webs are fairly common, despite being
less effective at intercepting and retaining prey and more vulnerable to damage by rain and falling debris.
Various researchers have suggested that horizontal webs offer compensating advantages, such as reduced
vulnerability to wind damage; reduced visibility to prey flying upwards, because of the backlighting from the sky;
enabling 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.
 Socialization
 A few spider species that build webs live together in large colonies and show social
behavior, although not as complex as in social insects. Anelosimus eximius (in the
family Theridiidae) can form colonies of up to 50,000 individuals. 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. In experiments, spider species like Steatoda grossa,
Latrodectus hesperus and Eratigena agrestis stayed away from Myrmica rubra ant
colonies. These ants are predators and pheromones they release for communication
have a notable deterrent effect on these spider species.