Arachnid

Arachnids are arthropods in the class Arachnida

(/əˈræknɪdə/) of the subphylum Chelicerata. Arachnid
Arachnida includes, among others, spiders, scorpions, Temporal range: Early Silurian – present
ticks, mites, pseudoscorpions, harvestmen, camel
spiders, whip spiders and vinegaroons.[2]

Adult arachnids have eight legs attached to the

cephalothorax. In some species the frontmost pair of
legs has converted to a sensory function, while in
others, different appendages can grow large enough
to take on the appearance of extra pairs of legs.

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 110,000 named species, of which 51,000 are
species of spiders.[3][4]
Representatives of the 12 extant orders of
The term is derived from the Greek word ἀράχνη
arachnids
(aráchnē, 'spider'), from the myth of the hubristic
human weaver Arachne, who was turned into a Scientific classification
spider.[5] Domain: Eukaryota
Kingdom: Animalia

Morphology Phylum: Arthropoda

Clade: Arachnomorpha
Subphylum: Chelicerata
Class: Arachnida
Lamarck, 1801

Orders

Ricinulei
?Xiphosura – horseshoe crabs
Basic characteristics of arachnids
include four pairs of legs (1) and a ?Eurypterida - sea scorpions
body divided into two tagmata: the Opiliones – harvestmen
cephalothorax (2) and the abdomen
(3) Solifugae – camel spiders
Acariformes – various mites
Almost all adult arachnids have eight legs, unlike Parasitiformes – various mites and
adult insects which all have six legs. However, ticks
arachnids also have two further pairs of appendages
†Phalangiotarbida (extinct)
that have become adapted for feeding, defense, and
sensory perception. The first pair, the chelicerae, Palpigradi – micro-whipscorpions
serve in feeding and defense. The next pair, the Arachnopulmonata
pedipalps, have been adapted for feeding,
locomotion, and/or reproductive functions. In Panscorpiones[1]
scorpions, pseudoscorpions, and ricinuleids the
Scorpiones – scorpions
pedipalps end in a pair of pinchers, while in whip
scorpions, Schizomida, Amblypygi, and most Pseudoscorpiones –
harvestmen, they are raptorial and used for prey pseudoscorpions
capture.[6] In Solifugae, the palps are quite leg-like,
Pantetrapulmonata
so that these animals appear to have ten legs. The
larvae of mites and Ricinulei have only six legs; a †Douglassarachne
fourth pair usually appears when they moult into
†Trigonotarbida (extinct)
nymphs. However, mites are variable: as well as
eight, there are adult mites with six or, like in Tetrapulmonata
Eriophyoidea, even four legs.[7][8] While the adult †Xenarachne
males in some members of Podapolipidae have six
legs, the adult females have only a single pair.[9] Schizotarsata

Arachnids are further distinguished from insects by †Haptopoda (extinct)

the fact they do not have antennae or wings. Their Pedipalpi
body is organized into two tagmata, called the
prosoma and opisthosoma, also referred to as the Amblypygi – whip
cephalothorax and abdomen. However, there are spiders
questions about the validity of the latter terms. While Schizomida – short-
the term cephalothorax implies a fused cephalon tailed whipscorpions
(head) and thorax, there is currently neither fossil nor
Uropygi –
embryological evidence that arachnids ever had a
separate thorax-like division. Likewise, the 'abdomen' vinegaroons
of many arachnids contains organs atypical of an Serikodiastida
abdomen, such as a heart and respiratory organs.[10]
†Idmonarachne
The cephalothorax is usually covered by a single,
†Uraraneida (extinct)
unsegmented carapace. The abdomen is segmented in
the more primitive forms, but varying degrees of Araneae – spiders
fusion 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 mites, the abdominal sections are completely fused.[11] A
telson is present in scorpions, where it has been modified to a stinger, and into a flagellum in the
Palpigradi, Schizomida (very short) and whip scorpions.[12] At the base of the flagellum in the two latter
groups there are glands which produce acetic acid as a chemical defense.[13] Except for a pair of pectines
in scorpions,[14] and the spinnerets in spiders, the abdomen has no appendages.[15]
Like all arthropods, arachnids have an exoskeleton, and they also have an internal structure of cartilage-
like tissue, called the endosternite, to which certain muscle groups are attached. The endosternite is even
calcified in some Opiliones.[16]

Locomotion
Most arachnids lack extensor muscles in the distal joints of their appendages. Spiders and whip scorpions
extend their limbs hydraulically using the pressure of their hemolymph.[17] Solifuges and some
harvestmen extend their knees by the use of highly elastic thickenings in the joint cuticle.[17] Scorpions,
pseudoscorpions and some harvestmen have evolved muscles that extend two leg joints (the femur-patella
and patella-tibia joints) at once.[18][19] The equivalent joints of the pedipalps of scorpions though, are
extended by elastic recoil.[20]

Physiology
There are characteristics that are particularly important for the
terrestrial lifestyle of arachnids, such as 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.[21] While the tracheae are often
individual systems of tubes, similar to those in insects, ricinuleids,
pseudoscorpions, and some 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 indicates that the
tracheae of arachnids are not homologous with those of insects.[22]

Further adaptations to terrestrial life are appendages modified for "Arachnida" from Ernst Haeckel's
more efficient locomotion on land, internal fertilisation, special Kunstformen der Natur, 1904
sensory organs, and water conservation enhanced by efficient
excretory structures as well as a waxy layer covering the cuticle.

The excretory glands of arachnids include up to four pairs of coxal glands along the side of the prosoma,
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
arachnids is guanine.[22]
Arachnid blood is variable in composition, depending on the mode of respiration. Arachnids with an
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-
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.[22]

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.[23] 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.[24][25][26] 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,[27] although it is not unusual for spiders to eat their own silk. And one species of spider is
mostly herbivorous.[28] Scorpions, spiders and pseudoscorpions secrete venom from specialized glands to
kill prey or defend themselves.[29] Their venom also contains pre-digestive enzymes that helps breaking
down the prey.[30][31][32] The saliva of ticks contains anticoagulants and anticomplements, and several
species produce a neurotoxin.[33][34]

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 pre-buccal cavity located immediately in front of the mouth. Behind the mouth is a
muscular, sclerotised pharynx, 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.[22]

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.[27] 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.[22]

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.[22]

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.[35] Complex
courtship rituals have evolved in many arachnids to
ensure the safe delivery of the sperm to the female.[22]
Members of many orders exhibit sexual dimorphism.[36]
Courtship behavior of Thelyphonus sp.
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.[37] In most arachnids only the females provide
parental care, with harvestmen being one of the few exceptions.[38][39]

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).[40][41][42][43][44] The three groups are related as shown in the cladogram below.[42] Including fossil
taxa does not fundamentally alter this view, although it introduces some additional basal groups.[45]

Chelicerata (sea spiders, horseshoe crabs and arachnids)

⁠
⁠

Arthropoda Myriapoda (centipedes, millipedes, and allies)

⁠ ⁠
⁠
⁠Mandibulata
⁠
⁠ Pancrustacea (crustaceans and hexapods)
⁠

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.[41][44] (Pycnogonida (sea spiders) may
be excluded from the chelicerates, which are then identified as the group labelled "Euchelicerata".[46]) A
2019 analysis nests Xiphosura deeply within Arachnida.[47]

⁠ ⁠ 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.[44]

⁠ Acariformes
⁠

⁠ ⁠ Opiliones
⁠ ⁠

⁠ ⁠ Ricinulei
⁠ ⁠
⁠
⁠
⁠ Solifugae
⁠

⁠ Parasitiformes
⁠

Arachnida ⁠
⁠ ⁠ Pseudoscorpiones
⁠

⁠
Arachnopulmonata
⁠ 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. Somewhat unexpectedly, there
was support for a clade comprising Opiliones, Ricinulei and
Solifugae, a combination not found in most other studies.[44] 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 Hubbardia pentapeltis (Schizomida)
considered may be due to long branch attraction.[47] 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,[48][49] and analyses support pseudoscorpions as the sister group of
scorpions.[50][51] Genetic analysis has not yet been done for Ricinulei, Palpigradi, or Solifugae, but
horseshoe crabs have gone through two whole genome duplications, which gives them five Hox clusters
with 34 Hox genes, the highest number found in any invertebrate, yet it is not clear if the oldest genome
duplication is related to the one in Arachnopulmonata.[52]

⁠ Panarthropoda
⁠ ⁠ Onychophora
⁠ ⁠
⁠
⁠ ⁠ Mandibulata
⁠ ⁠

⁠Chelicerata
⁠ Pycnogonida
⁠

⁠Euchelicerata
⁠ †Chasmataspidida
⁠

⁠Sclerophorata

⁠ †Eurypterida
⁠

⁠Arachnida ⁠ ⁠
⁠ ⁠ Parasitiformes
⁠ ⁠
⁠ Acariformes
⁠ ⁠
⁠
⁠
⁠
⁠ Pseudoscorpiones
⁠

⁠ ⁠ Opiliones
⁠ ⁠

⁠ Palpigradi
⁠
⁠
⁠
⁠ Solifugae
⁠
⁠
⁠ ⁠ ⁠
⁠ Ricinulei
⁠ ⁠
⁠
⁠ ⁠
⁠ Xiphosura
⁠ ⁠

⁠ ⁠
⁠

⁠
⁠
⁠
⁠

⁠Arachnopulmonata
⁠
⁠Pantetrapulmona
⁠
More recent phylogenomic analyses that have densely sampled both genomic datasets and morphology
have supported horseshoe crabs as nested inside Arachnida, suggesting a complex history of
terrestrialization.[53][54] Morphological analyses including fossils tend to recover the Tetrapulmonata,
including the extinct group the Haptopoda,[55][56][57][58][59] 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.[60]

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.[61][55] Fossil Goniotarbus angulatus
(Phalangiotarbida)

Taxonomy
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
Fossil of Kreischeria
are approximate. (Trigonotarbida)

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 (250 species)
Araneae – spiders (51,000 species)
Opiliones – phalangids, harvestmen or daddy-long-legs
(6,700 species)
Palpigradi – microwhip scorpions (130 species)
Parasitiformes – ticks (12,000 species) Eukoenenia spelaea (Palpigradi)

Pseudoscorpionida – pseudoscorpions (4,000 species)

Ricinulei – ricinuleids, hooded tickspiders (100 species)
Schizomida – "split middle" whip scorpions with divided exoskeletons (350 species)
Scorpiones – scorpions (2,700 species)
Solifugae – solpugids, windscorpions, sun spiders or camel spiders (1,200 species)
Uropygi (also called Thelyphonida) – whip scorpions or vinegaroons, forelegs modified into
sensory appendages and a long tail on abdomen tip (120 species)
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 110,000 arachnid species have been described, and that there may be over a million in
total.[4]

See also
Arthropods portal

Arachnophobia
Endangered spiders
Glossary of spider terms
List of extinct arachnids

References
1. Gainett, Guilherme; Klementz, Benjamin C.; Setton, Emily V. W.; Simian, Catalina; Iuri,
Hernán A.; Edgecombe, Gregory D.; Peretti, Alfredo V.; Sharma, Prashant P. (July 2024). "A
plurality of morphological characters need not equate with phylogenetic accuracy: A rare
genomic change refutes the placement of Solifugae and Pseudoscorpiones in
Haplocnemata" (https://onlinelibrary.wiley.com/doi/10.1111/ede.12467). Evolution &
Development. 26 (4). doi:10.1111/ede.12467 (https://doi.org/10.1111%2Fede.12467).
hdl:11086/552527 (https://hdl.handle.net/11086%2F552527). ISSN 1520-541X (https://searc
h.worldcat.org/issn/1520-541X).
2. Cracraft, Joel & Donoghue, Michael, eds. (2004). Assembling the Tree of Life (https://archiv
e.org/details/assemblingtreeli00crac_335). Oxford University Press. p. 297 (https://archive.o
rg/details/assemblingtreeli00crac_335/page/n314).
3. Brabazon, Anthony (2018). Foraging-Inspired Optimisation Algorithms. Springer
International Publishing. p. 237. ISBN 9783319591568.
4. Agnarsson, Ingi (2023). "Grand challenges in research on arachnid diversity, conservation,
and biogeography" (https://doi.org/10.3389%2Ffrchs.2023.1101141). Frontiers in Arachnid
Science. 2. doi:10.3389/frchs.2023.1101141 (https://doi.org/10.3389%2Ffrchs.2023.110114
1).
5. "Arachnid". Oxford English Dictionary (2nd ed.). 1989.
6. Schierwater, Bernd; DeSalle, Rob (2021-07-08). Invertebrate Zoology: A Tree of Life
Approach (https://books.google.com/books?id=Bk4vEAAAQBAJ&dq=%2522pincers+for+pr
ey+capture+in+scorpions%252C+pseudoscorpions%252C+and+Ricinulei%2522&pg=PA42
8). CRC Press. ISBN 978-1-4822-3582-1.
7. Schmidt, Günther (1993). Giftige und gefährliche Spinnentiere [Poisonous and dangerous
arachnids] (in German). Westarp Wissenschaften. p. 75. ISBN 978-3-89432-405-6.
8. Bolton, Samuel J.; Chetverikov, Philipp E.; Klompen, Hans (2017). "Morphological support
for a clade comprising two vermiform mite lineages: Eriophyoidea (Acariformes) and
Nematalycidae (Acariformes)" (https://doi.org/10.11158%2Fsaa.22.8.2). Systematic and
Applied Acarology. 22 (8): 1096. doi:10.11158/saa.22.8.2 (https://doi.org/10.11158%2Fsaa.
22.8.2). S2CID 90899467 (https://api.semanticscholar.org/CorpusID:90899467).
9. Dhooria, Manjit Singh (2016-12-14). Fundamentals of Applied Acarology (https://books.goog
le.com/books?id=ktS4DQAAQBAJ&dq=Podapolipidae+pair+of+legs&pg=PA49). Springer.
ISBN 978-981-10-1594-6.
10. Shultz, Stanley; Shultz, Marguerite (2009). The Tarantula Keeper's Guide. Hauppauge, New
York: Barron's. p. 23. ISBN 978-0-7641-3885-0.
11. Ruppert, E.; Fox, R. & Barnes, R. (2007). Invertebrate Zoology: A Functional Evolutionary
Approach (https://archive.org/details/isbn_9780030259821) (7th ed.). Thomson Learning.
ISBN 978-0-03-025982-1.
12. Little, Colin (1983-12-15). The Colonisation of Land: Origins and Adaptations of Terrestrial
Animals (https://books.google.com/books?id=tfs8AAAAIAAJ&dq=palpigrades+schizomida+t
elson&pg=PA108). Cambridge University Press. ISBN 978-0-521-25218-8.
13. Pinto-da-Rocha, Ricardo; Machado, Glauco; Giribet, Gonzalo (2007-02-28). Harvestmen:
The Biology of Opiliones (https://books.google.com/books?id=pbdpSKHkKDIC&dq=%2522A
mong+arachnids+%252C+chemical+defenses+have+been+recorded+for+Opiliones+%252
C+Uropygi+%252C+and+Schizomida%2522&pg=PA382). Harvard University Press.
ISBN 978-0-674-02343-7.
14. Di, Z.; Edgecombe, G. D.; Sharma, P. P. (2018). "Homeosis in a scorpion supports a
telopodal origin of pectines and components of the book lungs" (https://www.ncbi.nlm.nih.go
v/pmc/articles/PMC5963125). BMC Evolutionary Biology. 18 (1): 73.
Bibcode:2018BMCEE..18...73D (https://ui.adsabs.harvard.edu/abs/2018BMCEE..18...73D).
doi:10.1186/s12862-018-1188-z (https://doi.org/10.1186%2Fs12862-018-1188-z).
PMC 5963125 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5963125). PMID 29783957
(https://pubmed.ncbi.nlm.nih.gov/29783957).
15. Selden, Paul; Shear, William (2008). "Fossil evidence for the origin of spider spinnerets" (htt
ps://www.nature.com/articles/npre.2008.2088.1). Nature Precedings: 1.
doi:10.1038/npre.2008.2088.1 (https://doi.org/10.1038%2Fnpre.2008.2088.1).
16. Kovoor, J. (1978). "Natural calcification of the prosomatic endosternite in the Phalangiidae
(Arachnida:Opiliones)". Calcified Tissue Research. 26 (3): 267–269.
doi:10.1007/BF02013269 (https://doi.org/10.1007%2FBF02013269). PMID 750069 (https://p
ubmed.ncbi.nlm.nih.gov/750069). S2CID 23119386 (https://api.semanticscholar.org/CorpusI
D:23119386).
17. Sensenig, Andrew T. & Shultz, Jeffrey W. (February 15, 2003). "Mechanics of Cuticular
Elastic Energy Storage in Leg Joints Lacking Extensor Muscles in Arachnids". Journal of
Experimental Biology. 206 (4): 771–784. doi:10.1242/jeb.00182 (https://doi.org/10.1242%2F
jeb.00182). ISSN 1477-9145 (https://search.worldcat.org/issn/1477-9145). PMID 12517993
(https://pubmed.ncbi.nlm.nih.gov/12517993). S2CID 40503319 (https://api.semanticscholar.
org/CorpusID:40503319).
18. Shultz, Jeffrey W. (February 6, 2005). "Evolution of locomotion in arachnida: The hydraulic
pressure pump of the giant whipscorpion, Mastigoproctus giganteus (Uropygi)". Journal of
Morphology. 210 (1): 13–31. doi:10.1002/jmor.1052100103 (https://doi.org/10.1002%2Fjmo
r.1052100103). ISSN 1097-4687 (https://search.worldcat.org/issn/1097-4687).
PMID 29865543 (https://pubmed.ncbi.nlm.nih.gov/29865543). S2CID 46935000 (https://api.
semanticscholar.org/CorpusID:46935000).
19. Shultz, Jeffrey W. (January 1, 1992). "Muscle Firing Patterns in Two Arachnids Using
Different Methods of Propulsive Leg Extension" (http://jeb.biologists.org/content/162/1/313).
Journal of Experimental Biology. 162 (1): 313–329. doi:10.1242/jeb.162.1.313 (https://doi.or
g/10.1242%2Fjeb.162.1.313). ISSN 1477-9145 (https://search.worldcat.org/issn/1477-914
5). Retrieved 2012-05-19.
20. Sensenig, Andrew T. & Shultz, Jeffrey W. (2004). "Elastic energy storage in the pedipedal
joints of scorpions and sun-spiders (Arachnida, Scorpiones, Solifugae)" (https://www.biodive
rsitylibrary.org/part/228777). Journal of Arachnology. 32 (1): 1–10. doi:10.1636/S02-73 (http
s://doi.org/10.1636%2FS02-73). ISSN 0161-8202 (https://search.worldcat.org/issn/0161-820
2). S2CID 56461501 (https://api.semanticscholar.org/CorpusID:56461501).
21. Garwood, Russell J. & Edgecombe, Gregory D. (September 2011). "Early Terrestrial
Animals, Evolution, and Uncertainty" (https://doi.org/10.1007%2Fs12052-011-0357-y).
Evolution: Education and Outreach. 4 (3): 489–501. doi:10.1007/s12052-011-0357-y (http
s://doi.org/10.1007%2Fs12052-011-0357-y).
22. Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders
International. pp. 596–604. ISBN 978-0-03-056747-6.
23. Potapov, Anton M.; Beaulieu, Frédéric; Birkhofer, Klaus; Bluhm, Sarah L.; Degtyarev, Maxim
I.; Devetter, Miloslav; Goncharov, Anton A.; Gongalsky, Konstantin B.; Klarner, Bernhard;
Korobushkin, Daniil I.; Liebke, Dana F.; Maraun, Mark; Mc Donnell, Rory J.; Pollierer,
Melanie M.; Schaefer, Ina; Shrubovych, Julia; Semenyuk, Irina I.; Sendra, Alberto; Tuma,
Jiri; Tůmová, Michala; Vassilieva, Anna B.; Chen, Ting-Wen; Geisen, Stefan; Schmidt, Olaf;
Tiunov, Alexei V.; Scheu, Stefan (2022). "Feeding habits and multifunctional classification of
soil-associated consumers from protists to vertebrates" (https://doi.org/10.1111%2Fbrv.1283
2). Biological Reviews. 97 (3): 1057–1117. doi:10.1111/brv.12832 (https://doi.org/10.1111%
2Fbrv.12832). PMID 35060265 (https://pubmed.ncbi.nlm.nih.gov/35060265).
S2CID 246078291 (https://api.semanticscholar.org/CorpusID:246078291).
24. Powell, Erin C.; Painting, Christina J.; Hickey, Anthony J.; Machado, Glauco; Holwell,
Gregory I. (2021). "Diet, predators, and defensive behaviors of New Zealand harvestmen
(Opiliones: Neopilionidae)" (https://doi.org/10.1636%2FJoA-S-20-002). The Journal of
Arachnology. 49. doi:10.1636/JoA-S-20-002 (https://doi.org/10.1636%2FJoA-S-20-002).
S2CID 234364795 (https://api.semanticscholar.org/CorpusID:234364795).
25. Common harvestman | The Wildlife Trusts (https://www.wildlifetrusts.org/wildlife-explorer/inv
ertebrates/spiders/common-harvestman)
26. How do harvestmen hunt? - BBC Wildlife Magazine (https://www.discoverwildlife.com/anima
l-facts/insects-invertebrates/how-do-harvestmen-hunt/)
27. Machado, Glauco; Pinto-da-Rocha, Ricardo & Giribet, Gonzalo (2007). Pinto-da-Rocha,
Ricardo; Machado, Glauco & Giribet, Gonzalo (eds.). Harvestmen: the Biology of Opiliones.
Harvard University Press. ISBN 978-0-674-02343-7.
28. Rare Vegetarian Spider Discovered (https://www.livescience.com/5759-rare-vegetarian-spid
er-discovered.html)
29. Santibáñez-López, C. E.; Ontano, A. Z.; Harvey, M. S.; Sharma, P. P. (2018).
"Transcriptomic Analysis of Pseudoscorpion Venom Reveals a Unique Cocktail Dominated
by Enzymes and Protease Inhibitors" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC59832
63). Toxins. 10 (5): 207. doi:10.3390/toxins10050207 (https://doi.org/10.3390%2Ftoxins1005
0207). PMC 5983263 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5983263).
PMID 29783636 (https://pubmed.ncbi.nlm.nih.gov/29783636).
30. Zeh, J. A.; Bonilla, M. M.; Adrian, A. J.; Mesfin, S.; Zeh, D. W. (2012). "From father to son:
Transgenerational effect of tetracycline on sperm viability" (https://www.ncbi.nlm.nih.gov/pm
c/articles/PMC3337657). Scientific Reports. 2: 375. Bibcode:2012NatSR...2E.375Z (https://u
i.adsabs.harvard.edu/abs/2012NatSR...2E.375Z). doi:10.1038/srep00375 (https://doi.org/10.
1038%2Fsrep00375). PMC 3337657 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC33376
57). PMID 22540028 (https://pubmed.ncbi.nlm.nih.gov/22540028).
31. Delgado-Prudencio, G.; Cid-Uribe, J. I.; Morales, J. A.; Possani, L. D.; Ortiz, E.; Romero-
Gutiérrez, T. (2022). "The Enzymatic Core of Scorpion Venoms" (https://www.ncbi.nlm.nih.g
ov/pmc/articles/PMC9030722). Toxins. 14 (4): 248. doi:10.3390/toxins14040248 (https://doi.
org/10.3390%2Ftoxins14040248). PMC 9030722 (https://www.ncbi.nlm.nih.gov/pmc/article
s/PMC9030722). PMID 35448857 (https://pubmed.ncbi.nlm.nih.gov/35448857).
32. Walter, A.; Bechsgaard, J.; Scavenius, C.; Dyrlund, T. S.; Sanggaard, K. W.; Enghild, J. J.;
Bilde, T. (2017). "Characterisation of protein families in spider digestive fluids and their role
in extra-oral digestion" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5553785). BMC
Genomics. 18 (1): 600. doi:10.1186/s12864-017-3987-9 (https://doi.org/10.1186%2Fs12864
-017-3987-9). PMC 5553785 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5553785).
PMID 28797246 (https://pubmed.ncbi.nlm.nih.gov/28797246).
33. Denisov, S. S.; Ippel, J. H.; Castoldi, E.; Mans, B. J.; Hackeng, T. M.; Dijkgraaf, I. (2021).
"Molecular basis of anticoagulant and anticomplement activity of the tick salivary protein
Salp14 and its homologs" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8294578). The
Journal of Biological Chemistry. 297 (1): 100865. doi:10.1016/j.jbc.2021.100865 (https://doi.
org/10.1016%2Fj.jbc.2021.100865). PMC 8294578 (https://www.ncbi.nlm.nih.gov/pmc/articl
es/PMC8294578). PMID 34118237 (https://pubmed.ncbi.nlm.nih.gov/34118237).
34. Tick Paralysis - StatPearls - NCBI Bookshelf (https://www.ncbi.nlm.nih.gov/books/NBK4704
78/)
35. McLean, C. J.; Garwood, R. J.; Brassey, C. A. (2018). "Sexual dimorphism in the Arachnid
orders" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6225839). PeerJ. 6: e5751.
doi:10.7717/peerj.5751 (https://doi.org/10.7717%2Fpeerj.5751). PMC 6225839 (https://ww
w.ncbi.nlm.nih.gov/pmc/articles/PMC6225839). PMID 30416880 (https://pubmed.ncbi.nlm.ni
h.gov/30416880).
36. McLean, Callum J.; Garwood, Russell J.; Brassey, Charlotte A. (2018). "Sexual dimorphism
in the Arachnid orders" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6225839). PeerJ. 6:
e5751. doi:10.7717/peerj.5751 (https://doi.org/10.7717%2Fpeerj.5751). ISSN 2167-8359 (ht
tps://search.worldcat.org/issn/2167-8359). PMC 6225839 (https://www.ncbi.nlm.nih.gov/pm
c/articles/PMC6225839). PMID 30416880 (https://pubmed.ncbi.nlm.nih.gov/30416880).
37. Auerbach, Paul S. (2011-10-31). Wilderness Medicine E-Book: Expert Consult Premium
Edition - Enhanced Online Features (https://books.google.com/books?id=tdTInPqMCjMC&d
q=%2522some+are+ovoviviparous%252C+and+a+few+are+viviparous%2522&pg=PA950).
Elsevier Health Sciences. ISBN 978-1-4557-3356-9.
38. Quesada-Hidalgo, Rosannette; Solano-Brenes, Diego; Requena, Gustavo S.; Machado,
Glauco (April 2019). "The good fathers: efficiency of male care and the protective role of
foster parents in a Neotropical arachnid". Animal Behaviour. 150: 147–155.
doi:10.1016/j.anbehav.2019.02.007 (https://doi.org/10.1016%2Fj.anbehav.2019.02.007).
S2CID 73728615 (https://api.semanticscholar.org/CorpusID:73728615).
39. Nazareth, Tais M.; Machado, Glauco (March 2010). "Mating system and exclusive
postzygotic paternal care in a Neotropical harvestman (Arachnida: Opiliones)". Animal
Behaviour. 79 (3): 547–554. doi:10.1016/j.anbehav.2009.11.026 (https://doi.org/10.1016%2
Fj.anbehav.2009.11.026). S2CID 53166528 (https://api.semanticscholar.org/CorpusID:5316
6528).
40. Meusemann, Karen; Reumont, Björn M. von; Simon, Sabrina; Roeding, Falko; Strauss,
Sascha; Kück, Patrick; Ebersberger, Ingo; Walzl, Manfred; Pass, Günther; Breuers,
Sebastian; Achter, Viktor; Haeseler, Arndt von; Burmester, Thorsten; Hadrys, Heike; Wägele,
J. Wolfgang & Misof, Bernhard (2010). "A Phylogenomic Approach to Resolve the Arthropod
Tree of Life" (https://doi.org/10.1093%2Fmolbev%2Fmsq130). Molecular Biology and
Evolution. 27 (11): 2451–2464. doi:10.1093/molbev/msq130 (https://doi.org/10.1093%2Fmol
bev%2Fmsq130). PMID 20534705 (https://pubmed.ncbi.nlm.nih.gov/20534705).
41. Regier, Jerome C.; Shultz, Jeffrey W.; Zwick, Andreas; Hussey, April; Ball, Bernard; Wetzer,
Regina; Martin, Joel W. & Cunningham, Clifford W. (2010). "Arthropod relationships
revealed by phylogenomic analysis of nuclear protein-coding sequences". Nature. 463
(7284): 1079–1083. Bibcode:2010Natur.463.1079R (https://ui.adsabs.harvard.edu/abs/2010
Natur.463.1079R). doi:10.1038/nature08742 (https://doi.org/10.1038%2Fnature08742).
PMID 20147900 (https://pubmed.ncbi.nlm.nih.gov/20147900). S2CID 4427443 (https://api.s
emanticscholar.org/CorpusID:4427443).
42. Rota-Stabelli, Omar; Campbell, Lahcen; Brinkmann, Henner; Edgecombe, Gregory D.;
Longhorn, Stuart J.; Peterson, Kevin J.; Pisani, Davide; Philippe, Hervé & Telford,
Maximilian J. (2010). "A congruent solution to arthropod phylogeny: phylogenomics,
microRNAs and morphology support monophyletic Mandibulata" (https://www.ncbi.nlm.nih.g
ov/pmc/articles/PMC3013382). Proceedings of the Royal Society of London B: Biological
Sciences. 278 (1703): 298–306. doi:10.1098/rspb.2010.0590 (https://doi.org/10.1098%2Frs
pb.2010.0590). PMC 3013382 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3013382).
PMID 20702459 (https://pubmed.ncbi.nlm.nih.gov/20702459).
43. Campbell, Lahcen I.; Rota-Stabelli, Omar; Edgecombe, Gregory D.; Marchioro, Trevor;
Longhorn, Stuart J.; Telford, Maximilian J.; Philippe, Hervé; Rebecchi, Lorena; Peterson,
Kevin J. & Pisani, Davide (2011). "MicroRNAs and phylogenomics resolve the relationships
of Tardigrada and suggest that velvet worms are the sister group of Arthropoda" (https://ww
w.ncbi.nlm.nih.gov/pmc/articles/PMC3179045). Proceedings of the National Academy of
Sciences. 108 (38): 15920–15924. Bibcode:2011PNAS..10815920C (https://ui.adsabs.harva
rd.edu/abs/2011PNAS..10815920C). doi:10.1073/pnas.1105499108 (https://doi.org/10.107
3%2Fpnas.1105499108). PMC 3179045 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC31
79045). PMID 21896763 (https://pubmed.ncbi.nlm.nih.gov/21896763).
44. Sharma, Prashant P.; Kaluziak, Stefan T.; Pérez-Porro, Alicia R.; González, Vanessa L.;
Hormiga, Gustavo; Wheeler, Ward C. & Giribet, Gonzalo (2014-01-11). "Phylogenomic
Interrogation of Arachnida Reveals Systemic Conflicts in Phylogenetic Signal" (https://doi.or
g/10.1093%2Fmolbev%2Fmsu235). Molecular Biology and Evolution. 31 (11): 2963–2984.
doi:10.1093/molbev/msu235 (https://doi.org/10.1093%2Fmolbev%2Fmsu235).
PMID 25107551 (https://pubmed.ncbi.nlm.nih.gov/25107551).
45. Legg, David A.; Sutton, Mark D. & Edgecombe, Gregory D. (2013). "Arthropod fossil data
increase congruence of morphological and molecular phylogenies" (https://doi.org/10.103
8%2Fncomms3485). Nature Communications. 4: 2485. Bibcode:2013NatCo...4.2485L (http
s://ui.adsabs.harvard.edu/abs/2013NatCo...4.2485L). doi:10.1038/ncomms3485 (https://doi.
org/10.1038%2Fncomms3485). PMID 24077329 (https://pubmed.ncbi.nlm.nih.gov/2407732
9).
46. Giribet, Gonzalo; Edgecombe, Gregory D. & Wheeler, Ward C. (2001). "Arthropod
phylogeny based on eight molecular loci and morphology". Nature. 413 (6852): 157–161.
Bibcode:2001Natur.413..157G (https://ui.adsabs.harvard.edu/abs/2001Natur.413..157G).
doi:10.1038/35093097 (https://doi.org/10.1038%2F35093097). PMID 11557979 (https://pub
med.ncbi.nlm.nih.gov/11557979). S2CID 4431635 (https://api.semanticscholar.org/CorpusI
D:4431635).
47. Ballesteros, J.A.; Sharma, P.P. (2019). "A critical appraisal of the placement of Xiphosura
(Chelicerata) with account of known sources of phylogenetic error" (https://doi.org/10.1093%
2Fsysbio%2Fsyz011). Systematic Biology. 68 (6): 896–917. doi:10.1093/sysbio/syz011 (http
s://doi.org/10.1093%2Fsysbio%2Fsyz011). PMID 30917194 (https://pubmed.ncbi.nlm.nih.go
v/30917194).
48. Gainett, Guilherme; González, Vanessa L.; Ballesteros, Jesús A.; Setton, Emily V. W.;
Baker, Caitlin M.; Barolo Gargiulo, Leonardo; Santibáñez-López, Carlos E.; Coddington,
Jonathan A.; Sharma, Prashant P. (2021). "The genome of a daddy-long-legs (Opiliones)
illuminates the evolution of arachnid appendages" (https://www.ncbi.nlm.nih.gov/pmc/article
s/PMC8334856). Proceedings of the Royal Society B: Biological Sciences. 288 (1956).
bioRxiv 10.1101/2021.01.11.426205 (https://doi.org/10.1101%2F2021.01.11.426205).
doi:10.1098/rspb.2021.1168 (https://doi.org/10.1098%2Frspb.2021.1168). PMC 8334856 (ht
tps://www.ncbi.nlm.nih.gov/pmc/articles/PMC8334856). PMID 34344178 (https://pubmed.nc
bi.nlm.nih.gov/34344178).
49. Schwager, E.E.; Sharma, P.P.; Clark, T.; Leite, D.J.; Wierschin, T.; Pechmann, M.; Akiyama-
Oda, Y.; Esposito, L.; Bechsgaard, J.; Bilde, T.; Buffry, A.; Chao, H.; Dinh, H.; Doddapaneni,
H.V.; Dugan, S.; Eibner, C.; Extavour, C.; Funch, P.; Garb., J.; Gonzalez, L.B.; Gonzalez,
V.L.; Griffiths-Jones, S.; Han, Y.; Hayashi, C.; Hilbrant, M.; Hughes, D.S.T.; Janssen, R.;
Lee, S.L.; Maeso, I.; Murali, S.C.; Muzny, D.M.; da Fonseca, R.N.; Paese, C.L.B.; Qu, J.;
Ronshaugen, M.; Schomburg, C.; Schoenauer, A.; Stollewerk, A.; Torres-Oliva, M.; Turetzek,
N.; Vanthournout, B.; Werren, J.H.; Wolff, C.; Worley, K.C.; Bucher, G.; Gibbs, R.A.;
Coddington, J.; Oda, H.; Stanke, M.; Ayoub, N.A.; Prpic, N.M.; Flot, J.F.; Posnien, N.;
Richards, S.; McGregor, A.P. (July 2017). "The house spider genome reveals an ancient
whole-genome duplication during arachnid evolution" (https://www.ncbi.nlm.nih.gov/pmc/arti
cles/PMC5535294). BMC Biology. 15 (1): 62. doi:10.1186/s12915-017-0399-x (https://doi.or
g/10.1186%2Fs12915-017-0399-x). PMC 5535294 (https://www.ncbi.nlm.nih.gov/pmc/article
s/PMC5535294). PMID 28756775 (https://pubmed.ncbi.nlm.nih.gov/28756775).
50. Garbiec, Arnold; Christophoryová, Jana; Jędrzejowska, Izabela (2022). "Spectacular
alterations in the female reproductive system during the ovarian cycle and adaptations for
matrotrophy in chernetid pseudoscorpions (Pseudoscorpiones: Chernetidae)" (https://www.n
cbi.nlm.nih.gov/pmc/articles/PMC9018881). Scientific Reports. 12 (1): 6447.
Bibcode:2022NatSR..12.6447G (https://ui.adsabs.harvard.edu/abs/2022NatSR..12.6447G).
doi:10.1038/s41598-022-10283-z (https://doi.org/10.1038%2Fs41598-022-10283-z).
PMC 9018881 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC9018881). PMID 35440674
(https://pubmed.ncbi.nlm.nih.gov/35440674).
51. Ontano, A.Z.; Gainett, G.; Aharon, S.; Ballesteros, J.A.; Benavides, L.R.; Corbett, K.F.; et al.
(June 2021). "Taxonomic sampling and rare genomic changes overcome long-branch
attraction in the phylogenetic placement of Pseudoscorpions" (https://academic.oup.com/mb
e/article/38/6/2446/6132263?login=false). Molecular Biology and Evolution. 38 (6): 2446–
2467. doi:10.1093/molbev/msab038 (https://doi.org/10.1093%2Fmolbev%2Fmsab038).
PMC 8136511 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8136511). PMID 33565584
(https://pubmed.ncbi.nlm.nih.gov/33565584).
52. Shingate, Prashant; Ravi, Vydianathan; Prasad, Aravind; Tay, Boon-Hui; Garg, Kritika M.;
Chattopadhyay, Balaji; Yap, Laura-Marie; Rheindt, Frank E.; Venkatesh, Byrappa (2020).
"Chromosome-level assembly of the horseshoe crab genome provides insights into its
genome evolution" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7210998). Nature
Communications. 11 (1): 2322. Bibcode:2020NatCo..11.2322S (https://ui.adsabs.harvard.ed
u/abs/2020NatCo..11.2322S). doi:10.1038/s41467-020-16180-1 (https://doi.org/10.1038%2
Fs41467-020-16180-1). ISSN 2041-1723 (https://search.worldcat.org/issn/2041-1723).
PMC 7210998 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7210998). PMID 32385269
(https://pubmed.ncbi.nlm.nih.gov/32385269).
53. Ballesteros, Jesús A.; Santibáñez López, Carlos E.; Kováč, Ľubomír; Gavish-Regev, Efrat;
Sharma, Prashant P. (2019-12-18). "Ordered phylogenomic subsampling enables diagnosis
of systematic errors in the placement of the enigmatic arachnid order Palpigradi" (https://ww
w.ncbi.nlm.nih.gov/pmc/articles/PMC6939912). Proceedings of the Royal Society B:
Biological Sciences. 286 (1917): 20192426. doi:10.1098/rspb.2019.2426 (https://doi.org/10.
1098%2Frspb.2019.2426). ISSN 0962-8452 (https://search.worldcat.org/issn/0962-8452).
PMC 6939912 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6939912). PMID 31847768
(https://pubmed.ncbi.nlm.nih.gov/31847768).
54. Ballesteros, Jesús A.; Santibáñez-López, Carlos E.; Baker, Caitlin M.; Benavides, Ligia R.;
Cunha, Tauana J.; Gainett, Guilherme; et al. (2022-02-03). "Comprehensive species
sampling and sophisticated algorithmic approaches refute the monophyly of Arachnida" (http
s://academic.oup.com/mbe/article/doi/10.1093/molbev/msac021/6522129). Molecular
Biology and Evolution. 39 (2): msac021. doi:10.1093/molbev/msac021 (https://doi.org/10.10
93%2Fmolbev%2Fmsac021). ISSN 0737-4038 (https://search.worldcat.org/issn/0737-403
8). PMC 8845124 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8845124).
PMID 35137183 (https://pubmed.ncbi.nlm.nih.gov/35137183).
55. Wang, B.; Dunlop, J.A.; Selden, P.A.; Garwood, R.J.; Shear, W.A.; Müller, P.; Lei, X. (2018).
"Cretaceous arachnid Chimerarachne yingi gen. et sp. nov. illuminates spider origins" (http
s://www.research.manchester.ac.uk/portal/en/publications/cretaceous-arachnid-chimerarach
ne-yingi-gen-et-sp-nov-illuminates-spider-origins(b82d83bd-6081-4187-acfc-fb7358c33358).
html). Nature Ecology & Evolution. 2 (4): 614–622. Bibcode:2018NatEE...2..614W (https://ui.
adsabs.harvard.edu/abs/2018NatEE...2..614W). doi:10.1038/s41559-017-0449-3 (https://do
i.org/10.1038%2Fs41559-017-0449-3). PMID 29403075 (https://pubmed.ncbi.nlm.nih.gov/2
9403075). S2CID 4239867 (https://api.semanticscholar.org/CorpusID:4239867).
56. Garwood, R.J.; Dunlop, J.A.; Knecht, B.J.; Hegna, T.A. (2017). "The phylogeny of fossil whip
spiders" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5399839). BMC Evolutionary
Biology. 17 (1): 105. Bibcode:2017BMCEE..17..105G (https://ui.adsabs.harvard.edu/abs/20
17BMCEE..17..105G). doi:10.1186/s12862-017-0931-1 (https://doi.org/10.1186%2Fs12862-
017-0931-1). PMC 5399839 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5399839).
PMID 28431496 (https://pubmed.ncbi.nlm.nih.gov/28431496).
57. Garwood, R.J.; Dunlop, J.A.; Selden, P.A.; Spencer, A.R.T.; Atwood, R.C.; Vo, N.T.;
Drakopoulos, M. (2016). "Almost a spider: A 305 million-year-old fossil arachnid and spider
origins" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4822468). Proceedings of the Royal
Society B: Biological Sciences. 283 (1827): 20160125. doi:10.1098/rspb.2016.0125 (https://
doi.org/10.1098%2Frspb.2016.0125). PMC 4822468 (https://www.ncbi.nlm.nih.gov/pmc/artic
les/PMC4822468). PMID 27030415 (https://pubmed.ncbi.nlm.nih.gov/27030415).
58. Garwood, R.J.; Dunlop, J. (2014). "Three-dimensional reconstruction and the phylogeny of
extinct chelicerate orders" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4232842). PeerJ.
2: e641. doi:10.7717/peerj.641 (https://doi.org/10.7717%2Fpeerj.641). PMC 4232842 (http
s://www.ncbi.nlm.nih.gov/pmc/articles/PMC4232842). PMID 25405073 (https://pubmed.ncbi.
nlm.nih.gov/25405073).
59. Shultz, J.W. (2007). "A phylogenetic analysis of the arachnid orders based on morphological
characters" (https://doi.org/10.1111%2Fj.1096-3642.2007.00284.x). Zoological Journal of
the Linnean Society. 150 (2): 221–265. doi:10.1111/j.1096-3642.2007.00284.x (https://doi.or
g/10.1111%2Fj.1096-3642.2007.00284.x).
60. Selden, P.A.; Shear, W.A. & Sutton, M.D. (2008), "Fossil evidence for the origin of spider
spinnerets, and a proposed arachnid order", Proceedings of the National Academy of
Sciences, 105 (52): 20781–20785, Bibcode:2008PNAS..10520781S (https://ui.adsabs.harva
rd.edu/abs/2008PNAS..10520781S), doi:10.1073/pnas.0809174106 (https://doi.org/10.107
3%2Fpnas.0809174106), PMC 2634869 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC26
34869), PMID 19104044 (https://pubmed.ncbi.nlm.nih.gov/19104044)
61. Briggs, Helen (5 February 2018). " 'Extraordinary' fossil sheds light on origins of spiders" (htt
ps://www.bbc.co.uk/news/science-environment-42945813). BBC. Retrieved 9 June 2018.

External links
Arachnid (http://www.nhm.ac.uk/our-science/collections/zoology-collections/arachnid-collecti
ons.html), Natural History Museum, London

Retrieved from "https://en.wikipedia.org/w/index.php?title=Arachnid&oldid=1264996979"