Arachnid

Arachnida (/əˈræknɪdə/) is a class of joint-legged invertebrate animals (arthropods), in the

subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, Arachnids

pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaroons.[1] Temporal range: Early Silurian – present

Adult arachnids have eight legs attached to the cephalothorax, although the frontmost pair of legs in
some species has converted to a sensory function, while in other species, different appendages can grow
large enough to take on the appearance of extra pairs of legs. The term is derived from the Greek word
ἀράχνη (aráchnē, 'spider'), from the myth of the hubristic human weaver Arachne, who was turned into
a spider.[2]

Almost all extant arachnids are terrestrial, living mainly on land. However, some inhabit freshwater
environments and, with the exception of the pelagic zone, marine environments as well. They comprise
over 100,000 named species, of which 47,000 are species of spiders.[3]

Morphology
Basic characteristics of arachnids
include four pairs of legs (1) and a
body divided into two tagmata: the
cephalothorax (2) and the abdomen
(3)
pair.[7]
Almost all adult arachnids have eight legs, unlike adult insects
which all have six legs. However, arachnids also have two further
pairs of appendages that have become adapted for feeding, defense,
and sensory perception. The first pair, the chelicerae, serve in Representatives of the 12 extant orders of
feeding and defense. The next pair of appendages, the pedipalps, arachnids
have been adapted for feeding, locomotion, and/or reproductive
functions. In scorpions, pseudoscorpions, and ricinuleids the Scientific classification
pedipalps ends in a pair of pinchers, and in whip scorpions, Kingdom: Animalia
Schizomida, Amblypygi, and most harvestmen, they are raptorial
Clade: Bilateria
and used for prey capture.[4] In Solifugae, the palps are quite leg-
like, so that these animals appear to have ten legs. The larvae of Clade: Nephrozoa
mites and Ricinulei have only six legs; a fourth pair usually appears
when they moult into nymphs. However, mites are variable: as well (unranked): Protostomia
as eight, there are adult mites with six or, like in Eriophyoidea, even Superphylum: Ecdysozoa
four legs.[5][6] And while the adult males in some members of
(unranked): Panarthropoda
Podapolipidae have six legs, the adult females have only a single
(unranked): Tactopoda

Arachnids are further distinguished from insects by the fact they do not have antennae or wings. Their Phylum: Arthropoda
body is organized into two tagmata, called the prosoma, or cephalothorax, and the opisthosoma, or Clade: Arachnomorpha
abdomen. (However, there is currently neither fossil nor embryological evidence that arachnids ever
had a separate thorax-like division, so the validity of the term cephalothorax, which means a fused Subphylum: Chelicerata
cephalon, or head, and thorax, has been questioned. There are also arguments against use of 'abdomen', Class: Arachnida

as the opisthosoma of many arachnids contains organs atypical of an abdomen, such as a heart and
Lamarck, 1801
respiratory organs.[8]) The prosoma, or cephalothorax, is usually covered by a single, unsegmented
carapace. The abdomen is segmented in the more primitive forms, but varying degrees of fusion Orders
between the segments occur in many groups. It is typically divided into a preabdomen and
postabdomen, although this is only clearly visible in scorpions, and in some orders, such as the Acari, Ricinulei
the abdominal sections are completely fused.[9] A telson is present in scorpions, where it has been
Opiliones – harvestmen
modified to a stinger, and into a flagellum in the Palpigradi, Schizomida (very short) and whip
scorpions.[10] At the base of the flagellum in the two latter groups there are gland who produce acetic Solifugae – camel spiders
acid as a chemical defense.[11] Except for a pair of pectines in scorpions,[12] and the spinnerets in Acariformes mites
spiders, the abdomen has no appendages.[13]
Parasitiformes mites and ticks
Like all arthropods, arachnids have an exoskeleton, and they also have an internal structure of
†Phalangiotarbida (extinct)
cartilage-like tissue, called the endosternite, to which certain muscle groups are attached. The
endosternite is even calcified in some Opiliones.[14] Palpigradi – micro-whipscorpions
Arachnopulmonata
Locomotion
Panscorpiones
Most arachnids lack extensor muscles in the distal joints of their appendages. Spiders and Pseudoscorpiones –
whipscorpions extend their limbs hydraulically using the pressure of their hemolymph.[15] Solifuges
pseudoscorpions
and some harvestmen extend their knees by the use of highly elastic thickenings in the joint cuticle.[15]
Scorpions, pseudoscorpions and some harvestmen have evolved muscles that extend two leg joints (the Scorpiones – scorpions
femur-patella and patella-tibia joints) at once.[16][17] The equivalent joints of the pedipalps of scorpions
Pantetrapulmonata
though, are extended by elastic recoil.[18]
†Trigonotarbida (extinct)
Physiology Tetrapulmonata

There are characteristics that are particularly important for the terrestrial lifestyle of arachnids, such as Schizotarsata
internal respiratory surfaces in the form of tracheae, or modification of the book gill into a book lung,
an internal series of vascular lamellae used for gas exchange with the air.[19] While the tracheae are †Haptopoda (extinct)
often individual systems of tubes, similar to those in insects, ricinuleids, pseudoscorpions, and some Pedipalpi
spiders possess sieve tracheae, in which several tubes arise in a bundle from a small chamber connected
to the spiracle. This type of tracheal system has almost certainly evolved from the book lungs, and Amblypygi – whip
indicates that the tracheae of arachnids are not homologous with those of insects.[20] spiders
Further adaptations to terrestrial life are appendages modified for more efficient locomotion on land, Schizomida – short-
internal fertilisation, special sensory organs, and water conservation enhanced by efficient excretory
tailed whipscorpions
structures as well as a waxy layer covering the cuticle.
Uropygi –
The excretory glands of arachnids include up to four pairs of coxal glands along the side of the prosoma, vinegaroons
and one or two pairs of Malpighian tubules, emptying into the gut. Many arachnids have only one or the
other type of excretory gland, although several do have both. The primary nitrogenous waste product in Serikodiastida
arachnids is guanine.[20]
†Uraraneida (extinct)
Arachnid blood is variable in composition, depending on the mode of respiration. Arachnids with an
Araneae – spiders
efficient tracheal system do not need to transport oxygen in the blood, and may have a reduced
circulatory system. In scorpions and some spiders, however, the blood contains haemocyanin, a copper- ?Xiphosura – horseshoe crabs
based pigment with a similar function to haemoglobin in vertebrates. The heart is located in the
forward part of the abdomen, and may or may not be segmented. Some mites have no heart at all.[20]

Diet and digestive system

Arachnids are mostly carnivorous, feeding on the pre-digested bodies of insects and other small animals. But ticks,
and many mites, are parasites, some of which are carriers of disease. The diet of mites also include tiny animals,
fungi, plant juices and decomposing matter.[21] Almost as varied is the diet of harvestmen, where we will find
predators, decomposers and omnivores feeding on decaying plant and animal matter, droppings, animals and
mushrooms.[22][23][24] The harvestmen and some mites, such as the house dust mite, are also the only arachnids
able to ingest solid food, which exposes them to internal parasites,[25] although it is not unusual for spiders to eat
their own silk. And one species of spider is mostly herbivorous.[26] Scorpions, spiders and pseudoscorpions secrete
venom from specialized glands to kill prey or defend themselves.[27] Their venom also contains pre-digestive
enzymes that helps breaking down the prey.[28][29][30] The saliva of ticks contains anticoagulants and
anticomplements, and several species produce a neurotoxin.[31][32]

Arachnids produce digestive enzymes in their stomachs, and use their pedipalps and chelicerae to pour them over
their dead prey. The digestive juices rapidly turn the prey into a broth of nutrients, which the arachnid sucks into a "Arachnida" from Ernst Haeckel's
pre-buccal cavity located immediately in front of the mouth. Behind the mouth is a muscular, sclerotised pharynx, Kunstformen der Natur, 1904
which acts as a pump, sucking the food through the mouth and on into the oesophagus and stomach. In some
arachnids, the oesophagus also acts as an additional pump.

The stomach is tubular in shape, with multiple diverticula extending throughout the body. The stomach and its diverticula both produce digestive
enzymes and absorb nutrients from the food. It extends through most of the body, and connects to a short sclerotised intestine and anus in the hind
part of the abdomen.[20]

Senses
Arachnids have two kinds of eyes: the lateral and median ocelli. The lateral ocelli evolved from compound eyes and may have a tapetum, which
enhances the ability to collect light. With the exception of scorpions, which can have up to five pairs of lateral ocelli, there are never more than three
pairs present. The median ocelli develop from a transverse fold of the ectoderm. The ancestors of modern arachnids probably had both types, but
modern ones often lack one type or the other.[25] The cornea of the eye also acts as a lens, and is continuous with the cuticle of the body. Beneath this
is a transparent vitreous body, and then the retina and, if present, the tapetum. In most arachnids, the retina probably does not have enough light
sensitive cells to allow the eyes to form a proper image.[20]

In addition to the eyes, almost all arachnids have two other types of sensory organs. The most important to most arachnids are the fine sensory hairs
that cover the body and give the animal its sense of touch. These can be relatively simple, but many arachnids also possess more complex structures,
called trichobothria.

Finally, slit sense organs are slit-like pits covered with a thin membrane. Inside the pit, a small hair touches the underside of the membrane, and
detects its motion. Slit sense organs are believed to be involved in proprioception, and possibly also hearing.[20]

Reproduction
Arachnids may have one or two gonads, which are located in the abdomen. The genital opening is usually
located on the underside of the second abdominal segment. In most species, the male transfers sperm to
the female in a package, or spermatophore. The males in harvestmen and some mites have a penis.[33]
Complex courtship rituals have evolved in many arachnids to ensure the safe delivery of the sperm to the
female.[20] Members of many orders exhibit sexual dimorphism.[34]

Arachnids usually lay yolky eggs, which hatch into immatures that resemble adults. Scorpions, however,
are either ovoviviparous or viviparous, depending on species, and bear live young. Also some mites are
ovoviviparous and viviparous, even if most lay eggs.[35] In most arachnids only the females provide
parental care, with harvestmen being one of the few exceptions.[36][37]
Courtship behavior of Thelyphonus sp.

Taxonomy and evolution

Phylogeny

The phylogenetic relationships among the main subdivisions of arthropods have been the subject of considerable research and dispute for many years.
A consensus emerged from about 2010 onwards, based on both morphological and molecular evidence. Extant (living) arthropods are a monophyletic
group and are divided into three main clades: chelicerates (including arachnids), pancrustaceans (the paraphyletic crustaceans plus insects and their
allies), and myriapods (centipedes, millipedes and allies).[38][39][40][41][42] The three groups are related as shown in the cladogram below.[40]
Including fossil taxa does not fundamentally alter this view, although it introduces some additional basal groups.[43]

⁠Arthropoda ⁠
 Chelicerata (sea spiders, horseshoe crabs and arachnids)

⁠Pancrustacea (crustaceans and hexapods)

⁠Mandibulata
⁠
Myriapoda (centipedes, millipedes, and allies)
⁠

The extant chelicerates comprise two marine groups: sea spiders and horseshoe crabs, and the terrestrial arachnids. These have been thought to be
related as shown below.[39][42] (Pycnogonida (sea spiders) may be excluded from the chelicerates, which are then identified as the group labelled
"Euchelicerata".[44]) A 2019 analysis nests Xiphosura deeply within Arachnida.[45]

⁠ Pycnogonida (sea spiders)

⁠Chelicerata ⁠Xiphosura (horseshoe crabs)

⁠Euchelicerata
⁠
Arachnida
⁠

Discovering relationships within the arachnids has proven difficult as of March 2016, with successive studies producing different results. A study in
2014, based on the largest set of molecular data to date, concluded that there were systematic conflicts in the phylogenetic information, particularly
affecting the orders Acariformes, Parasitiformes and Pseudoscorpiones, which have had much faster evolutionary rates. Analyses of the data using
sets of genes with different evolutionary rates produced mutually incompatible phylogenetic trees. The authors favoured relationships shown by more
slowly evolving genes, which demonstrated the monophyly of Chelicerata, Euchelicerata and Arachnida, as well as of some clades within the
arachnids. The diagram below summarizes their conclusions, based largely on the 200 most slowly evolving genes; dashed lines represent uncertain
placements.[42]

⁠Arachnida
⁠
Acariformes

⁠
Opiliones

⁠Ricinulei

⁠
Solifugae
⁠

Parasitiformes
Arachnopulmonata

⁠
Pseudoscorpiones

Scorpiones

⁠
Araneae

⁠
⁠
⁠

⁠Tetrapulmonata Amblypygi
⁠
 Uropygi (Thelyphonida s.s.)
⁠

Tetrapulmonata, here consisting of Araneae, Amblypygi and Uropygi (Thelyphonida s.s.) (Schizomida was not
included in the study), received strong support. The addition of Scorpiones to produce a clade called
Arachnopulmonata was also well supported. Pseudoscorpiones may also belong here, as all six orders share the
same ancient whole genome duplication,[46] and analyses support pseudoscorpions as the sister group of
scorpions.[47] Somewhat unexpectedly, there was support for a clade comprising Opiliones, Ricinulei and
Solifugae, a combination not found in most other studies.[42]
In early 2019, a molecular phylogenetic analysis
placed the horseshoe crabs, Xiphosura, as the sister group to Ricinulei. It also grouped pseudoscorpions with mites
and ticks, which the authors considered may be due to long branch attraction.[45] Hubbardia pentapeltis (Schizomida)

⁠Onychophora

⁠ ⁠
Mandibulata

⁠Chelicerata
⁠
⁠ Pycnogonida

⁠Euchelicerata
⁠
†Chasmataspidida
⁠

⁠Sclerophorata

†Eurypterida
⁠

⁠Arachnida
⁠Parasitiformes

⁠ ⁠Acariformes

⁠
Pseudoscorpiones
⁠

⁠
⁠Opiliones

⁠
⁠Palpigradi

Solifugae
⁠
⁠

⁠
Ricinulei

⁠
Xiphosura
⁠

⁠Arachnopulmonata
⁠
⁠ Scorpiones

⁠Pantetrapulmonata ⁠
⁠
 †Trigonotarbida

†Haptopoda
⁠

⁠Thelyphonid
⁠Schizotarsata

⁠Pedipalpi
⁠Tetrapulmonata
⁠

⁠
†Uraraneida

⁠Serikodiastida
⁠
Araneae
⁠

More recent phylogenomic analyses that have densely sampled both genomic datasets and morphology have consistently supported horseshoe crabs
as nested inside Arachnida, suggesting a complex history of terrestrialization.[48][49] Morphological analyses including fossils tend to recover the
Tetrapulmonata, including the extinct group the Haptopoda,[50][51][52][53][54] but recover other ordinal relationships with low support.

Fossil history

The Uraraneida are an extinct order of spider-like arachnids from the Devonian and Permian.[55]

A fossil arachnid in 100 million year old (mya) amber from Myanmar, Chimerarachne yingi, has spinnerets (to
produce silk); it also has a tail, like the Palaeozoic Uraraneida, some 200 million years after other known fossils
with tails. The fossil resembles the most primitive living spiders, the mesotheles.[56][50]

Taxonomy Fossil Goniotarbus angulatus

(Phalangiotarbida)
The subdivisions of the arachnids are usually treated as orders. Historically, mites and ticks were treated as a
single order, Acari. However, molecular phylogenetic studies suggest that the two groups do not form a single
clade, with morphological similarities being due to convergence. They are now usually treated as two separate taxa
– Acariformes, mites, and Parasitiformes, ticks – which may be ranked as orders or superorders. The arachnid
subdivisions are listed below alphabetically; numbers of species are approximate.

Extant forms

Acariformes – mites (32,000 species)

Amblypygi – "blunt rump" tail-less whip scorpions with front legs modified into whip-like sensory structures as
long as 25 cm or more (153 species)
Araneae – spiders (40,000 species) Fossil of Kreischeria
(Trigonotarbida)
Opiliones – phalangids, harvestmen or daddy-long-legs (6,300 species)
Palpigradi – microwhip scorpions (80 species)
Parasitiformes – ticks (12,000 species)
Pseudoscorpionida – pseudoscorpions (3,000 species)
Ricinulei – ricinuleids, hooded tickspiders (60 species)
Schizomida – "split middle" whip scorpions with divided exoskeletons (220 species)
Scorpiones – scorpions (2,000 species)
Solifugae – solpugids, windscorpions, sun spiders or camel spiders (900 species)
Uropygi (also called Thelyphonida) – whip scorpions or vinegaroons, forelegs modified into sensory
appendages and a long tail on abdomen tip (100 species)
Eukoenenia spelaea (Palpigradi)
Extinct forms

†Haptopoda – extinct arachnids apparently part of the Tetrapulmonata, the group including spiders and whip scorpions (1 species)
†Phalangiotarbida – extinct arachnids of uncertain affinity (30 species)
†Trigonotarbida – extinct (late Silurian Early Permian)
†Uraraneida – extinct spider-like arachnids, but with a "tail" and no spinnerets (2 species)

It is estimated that 98,000 arachnid species have been described, and that there may be up to 600,000 in total.[57]

See also
 Arthropods portal
Arachnophobia
Endangered spiders
Glossary of spider terms
List of extinct arachnids
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External links
Arachnid (http://www.nhm.ac.uk/our-science/collections/zoology-collections/arachnid-collections.html), Natural History Museum, London

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