Arachnid

Last updated

Arachnids
Temporal range: 430–0  Ma
Haeckel Arachnida.jpg
"Arachnida" from Ernst Haeckel's Kunstformen der Natur , 1904
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Lamarck, 1801
Orders

Arachnida ( /əˈræknɪdə/ ) is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Spiders are the largest order in the class, which also includes scorpions, ticks, mites, harvestmen, and solifuges. [1] In 2019, a molecular phylogenetic study also placed horseshoe crabs in Arachnida. [2]

Contents

Almost all adult arachnids have eight legs, although the front 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), from the myth of the hubristic human weaver Arachne, who was turned into a spider. [3]

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.

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) Spider-characteristics.png
Basic characteristics of arachnids include four pairs of legs (1) and a body divided into two tagmata: the cephalothorax (2) and the abdomen (3)

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 feeding and defense. The next pair of appendages, the pedipalps, have been adapted for feeding, locomotion, and/or reproductive functions. In Solifugae, the palps are quite leg-like, so that these animals appear to have ten legs. The larvae of mites and Ricinulei have only six legs; a fourth pair usually appears when they moult into nymphs. However, mites are variable: as well as eight, there are adult mites with six or even four legs. [4]

Arachnids are further distinguished from insects by the fact they do not have antennae or wings. Their body is organized into two tagmata, called the prosoma, or cephalothorax, and the opisthosoma, or 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 cephalon, or head, and thorax, has been questioned. There are also arguments against use of 'abdomen', as the opisthosoma of many arachnids contains organs atypical of an abdomen, such as a heart and respiratory organs. [5] ) 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 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, the abdominal sections are completely fused. [6] A telson is present in scorpions, where it has been modified to a stinger, and in the Schizomida, whip scorpions and Palpigradi. [7]

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. [8]

Locomotion

Most arachnids lack extensor muscles in the distal joints of their appendages. Spiders and whipscorpions extend their limbs hydraulically using the pressure of their hemolymph. [9] Solifuges and some harvestmen extend their knees by the use of highly elastic thickenings in the joint cuticle. [9] Scorpions, pseudoscorpions and some harvestmen have evolved muscles that extend two leg joints (the femur-patella and patella-tibia joints) at once. [10] [11] The equivalent joints of the pedipalps of scorpions though, are extended by elastic recoil. [12]

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. [13] 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. [14]

Further adaptations to terrestrial life are appendages modified for more efficient locomotion on land, internal fertilisation, special 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. [14]

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. [14]

Diet and digestive system

Arachnids are mostly carnivorous, feeding on the pre-digested bodies of insects and other small animals. Only in the harvestmen and among mites, such as the house dust mite, is there ingestion of solid food particles, and thus exposure to internal parasites, [15] although it is not unusual for spiders to eat their own silk. Several groups secrete venom from specialized glands to kill prey or enemies. Several mites and ticks are parasites, some of which are carriers of disease.

Arachnids produce digestive juices 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. [14]

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. [15] 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. [14]

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. [14]

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. Complex courtship rituals have evolved in many arachnids to ensure the safe delivery of the sperm to the female. [14] Members of many orders exhibit sexual dimorphism. [16]

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. In most arachnids only the females provide parental care, with harvestmen being one of the few exceptions.[ citation needed ]

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). [17] [18] [19] [20] [21] The three groups are related as shown in the cladogram below. [19] Including fossil taxa does not fundamentally alter this view, although it introduces some additional basal groups. [22]

Arthropoda

Chelicerata (sea spiders, horseshoe crabs and arachnids)

Mandibulata

Pancrustacea (crustaceans and insects)

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. [18] [21] (Pycnogonida (sea spiders) may be excluded from the chelicerates, which are then identified as the group labelled "Euchelicerata". [23] ) A 2019 analysis nests Xiphosura deeply within Arachnida. [2]

Chelicerata

Pycnogonida (sea spiders)

Euchelicerata

Xiphosura (horseshoe crabs)

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. [21]

Arachnida

Acariformes Trombidium holosericeum (aka).jpg

Opiliones Harvestman opilio canestrinii male.jpg

Ricinulei Cryptocellus goodnighti.jpg

Solifugae Galeodes.jpg

Parasitiformes

Pseudoscorpiones Ar 1.jpg

Scorpiones SCORPIO MAURUS PALMATUS.jpg

Tetrapulmonata

Araneae Araneus diadematus (aka).jpg

Amblypygi Amblypigid.jpg

Thelyphonida (Uropygi) Whipscorpion.jpg

Arachnopulmonata
Hubbardia pentapeltis (Schizomida) Hubbardia pentapeltis female.jpg
Hubbardia pentapeltis (Schizomida)

Tetrapulmonata, here consisting of Araneae, Amblypygi and Thelyphonida (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, possibly as the sister of Scorpiones. Somewhat unexpectedly, there was support for a clade comprising Opiliones, Ricinulei and Solifugae, a combination not found in most other studies. [21]

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. [2]

Onychophora

Mandibulata

Chelicerata

Pycnogonida

Euchelicerata

Parasitiformes

Acariformes

Pseudoscorpiones

Opiliones

Solifugae

Ricinulei

Xiphosura

Scorpiones

Tetrapulmonata

Morphological analyses including fossils tend to recover the Tetrapulmonata, including the extinct group the Haptopoda, [24] [25] [26] [27] [28] but recover other ordinal relationships with low support.

Fossil history

Fossil Goniotarbus angulatus (Phalangiotarbi) Goniotarbus angulatus holotype fossil dorsal ventral.jpg
Fossil Goniotarbus angulatus (Phalangiotarbi)
Fossil of Kreischeria (Trigonotarbida) Kreischeria Vienna.jpg
Fossil of Kreischeria (Trigonotarbida)

The Uraraneida are an extinct order of spider-like arachnids from the Devonian and Permian. [29]

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. [30] [24]

Taxonomy

Eukoenenia spelaea (Palpigradi) Live Eukoenenia spelaea in its cave habitat.png
Eukoenenia spelaea (Palpigradi)

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.

It is estimated that 98,000 arachnid species have been described, and that there may be up to 600,000 in total. [31]

See also

Related Research Articles

Chelicerata subphylum of arthropods

The subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda. It contains the sea spiders, arachnids, and several extinct lineages, such as the eurypterids.

Amblypygi order of arachnids

Amblypygi is an ancient order of arachnid chelicerate arthropods also known as whip spiders and tailless whip scorpions. The name "amblypygid" means "blunt tail", a reference to a lack of the flagellum that is otherwise seen in whip scorpions. They are harmless to humans. Amblypygids possess no silk glands or venomous fangs. They rarely bite if threatened, but can grab fingers with their pedipalps, resulting in thorn-like puncture injuries.

Opiliones Order of arachnids (harvestmen/daddy longlegs)

The Opiliones are an order of arachnids colloquially known as harvestmen, harvesters, or daddy longlegs. As of April 2017, over 6,650 species of harvestmen have been discovered worldwide, although the total number of extant species may exceed 10,000. The order Opiliones includes five suborders: Cyphophthalmi, Eupnoi, Dyspnoi, Laniatores, and Tetrophthalmi, which were named in 2014.

Thelyphonida order of arachnids

Thelyphonida is an arachnid order comprising invertebrates commonly known as whip scorpions or vinegaroons. They are often called uropygids in the scientific community based on an alternative name for the order, Uropygi. The name "whip scorpion" refers to their resemblance to true scorpions and possession of a whiplike tail. "Vinegaroon" is based on their ability when attacked to discharge an offensive liquid which contains acetic acid, producing a vinegar-like smell.

Ricinulei order of arachnids

The order Ricinulei is a group of arachnids known as hooded tickspiders, though they are not true spiders. Like most arachnids, they are predatory, eating small arthropods. In older works they are sometimes referred to as Podogona.

Book lung A type of lung commonly found in arachnids

A book lung is a type of respiration organ used for atmospheric gas exchange that is present in many arachnids, such as scorpions and spiders. Each of these organs is located inside an open ventral abdominal, air-filled cavity (atrium) and connects with the surroundings through a small opening for the purpose of respiration.

Acari Subclass of arachnids

The Acari are a taxon of arachnids that contains mites and ticks. The diversity of the Acari is extraordinary and their fossil history goes back to at least the early Devonian period. Acarologists have proposed a complex set of taxonomic ranks to classify mites. In most modern treatments, the Acari are considered a subclass of the Arachnida and are composed of two or three superorders or orders: Acariformes, Parasitiformes, and Opilioacariformes; the latter is often considered a subgroup within the Parasitiformes. The monophyly of the Acari is open to debate, and the relationships of the acarines to other arachnids is not at all clear. In older treatments, the subgroups of the Acarina were placed at order rank, but as their own subdivisions have become better understood, treating them at the superorder rank is more usual.

Pedipalp Appendage of chelicerate

Pedipalps are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") and anterior to the first pair of walking legs.

Sea spider Marine arthropod of class Pycnogonida

Sea spiders are marine arthropods of order Pantopoda, belonging to class Pycnogonida, hence they are also called pycnogonids. They are cosmopolitan, found in oceans around the world. There are over 1300 known species, with legs ranging from 1 mm (0.04 in) to over 70 cm (2.3 ft). Most are toward the smaller end of this range in relatively shallow depths; however, they can grow to be quite large in Antarctic and deep waters.

Solifugae order of animals in the class Arachnida

Solifugae is an order of animals in the class Arachnida known variously as camel spiders, wind scorpions, sun spiders, or solifuges. The order includes more than 1,000 described species in about 153 genera. Despite the common names, they are neither true scorpions nor true spiders. Most species of Solifugae live in dry climates and feed opportunistically on ground-dwelling arthropods and other small animals. The largest species grow to a length of 12–15 cm (5–6 in), including legs. A number of urban legends exaggerate the size and speed of the Solifugae, and their potential danger to humans, which is negligible.

<i>Plesiosiro</i> extinct arachnid species

Plesiosiro is an extinct arachnid genus known exclusively from only nine specimens from the Upper Carboniferous of Coseley, Staffordshire, United Kingdom. The genus is monotypic, represented only by the species Plesiosiro madeleyi described by Reginald Innes Pocock in his important 1911 monograph on British Carboniferous arachnids. It is the only known member of the order Haptopoda. The original locality from which these fossils originate is no longer available thus it is unclear whether any further examples will be found.

Tetrapulmonata

Tetrapulmonata is a non-ranked supra-ordinal clade of arachnids. It is composed of the extant orders Thelyphonida, Schizomida, Amblypygi and Araneae (spiders). It is the only supra-ordinal group of arachnids that is strongly supported in molecular phylogenetic studies. Two extinct orders are also placed in this clade, Haptopoda and Uraraneida. In 2016, a newly described fossil arachnid, Idmonarachne, was also included in the Tetrapulmonata; as of March 2016 it has not been assigned to an order.

Opisthosoma Wikimedia disambiguation page

The opisthosoma is the posterior part of the body in some arthropods, behind the prosoma (cephalothorax). It is a distinctive feature of the subphylum Chelicerata (arachnids, horseshoe crabs and others. Although it is similar in most respects to an abdomen, the opisthosoma is differentiated by its inclusion of the respiratory organs and the heart.

Evolution of spiders The origin from a chelicerate ancestor and diversification of spiders through geologic time

The evolution of spiders has been ongoing for at least 380 million years. The group's origins lie within an arachnid sub-group defined by the presence of book lungs ; the arachnids as a whole evolved from aquatic chelicerate ancestors. More than 45,000 extant species have been described, organised taxonomically in 3,958 genera and 114 families. There may be more than 120,000 species. Fossil diversity rates make up a larger proportion than extant diversity would suggest with 1,593 arachnid species described out of 1,952 recognized chelicerates. Both extant and fossil species are described yearly by researchers in the field. Major developments in spider evolution include the development of spinnerets and silk secretion.

Cyphophthalmi suborder of arachnids

Cyphophthalmi is a suborder of harvestmen, colloquially known as mite harvestmen. Cyphophthalmi comprises 36 genera, and more than two hundred described species. The six families are currently grouped into three infraorders: the Boreophthalmi, Scopulophthalmi, and Sternophthalmi.

Dromopoda subclass of arachnids

Dromopoda is a subclass of the arachnids, including the Opiliones (harvestmen), Scorpions, Pseudoscorpions and Solifugae. The latter three are sometimes grouped as Novogenuata. Combined morphological and molecular analyses have shown Dromopoda to be monophyletic. However, a strictly molecular analysis did not support the monophyly of Dromopoda.

Harvestman phylogeny

Harvestmen (Opiliones) are an order of arachnids often confused with spiders, though the two orders are not closely related. Research on harvestman phylogeny is in a state of flux. While some families are clearly monophyletic, that is share a common ancestor, others are not, and the relationships between families are often not well understood.

Arthropod Phylum of invertebrates with jointed exoskeletons

An arthropod is an invertebrate animal having an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Euarthropoda, which includes insects, arachnids, myriapods, and crustaceans. The term Arthropoda as originally proposed refers to a proposed grouping of Euarthropods and the phylum Onychophora.

Tetrophthalmi suborder of arachnids

Tetrophthalmi is an extinct suborder of Opiliones that had both median and lateral eyes. First described in 2014, it is known from two extinct species. Phylogenetic analysis suggests that this eye arrangement is the ancestral condition for harvestmen, placing Tetrophthalmi and Cyphophthalmi in a basal position within Opiliones.

References

  1. Cracraft, Joel & Donoghue, Michael, eds. (2004). Assembling the Tree of Life . Oxford University Press. p.  297.
  2. 1 2 3 4 Ballesteros, J. A.; Sharma, P. P. (2019). "A Critical Appraisal of the Placement of Xiphosura (Chelicerata) with Account of Known Sources of Phylogenetic Error". Systematic Biology. 68 (6): 896–917. doi: 10.1093/sysbio/syz011 . PMID   30917194.
  3. "Arachnid". Oxford English Dictionary (2nd ed.). 1989.
  4. Schmidt, Günther (1993). Giftige und gefährliche Spinnentiere[Poisonous and dangerous arachnids] (in German). Westarp Wissenschaften. p. 75. ISBN   978-3-89432-405-6.
  5. Shultz, Stanley; Shultz, Marguerite (2009). The Tarantula Keeper's Guide. Hauppauge, New York: Barron's. p. 23. ISBN   978-0-7641-3885-0.
  6. Ruppert, E.; Fox, R. & Barnes, R. (2007). Invertebrate Zoology: A Functional Evolutionary Approach (7th ed.). Thomson Learning. ISBN   978-0-03-025982-1.
  7. The Colonisation of Land: Origins and Adaptations of Terrestrial Animals
  8. 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. PMID   750069. S2CID   23119386.
  9. 1 2 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 . ISSN   1477-9145. PMID   12517993.
  10. 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. ISSN   1097-4687. PMID   29865543. S2CID   46935000.
  11. Shultz, Jeffrey W. (January 1, 1992). "Muscle Firing Patterns in Two Arachnids Using Different Methods of Propulsive Leg Extension". Journal of Experimental Biology. 162 (1): 313–329. ISSN   1477-9145 . Retrieved 2012-05-19.
  12. Sensenig, Andrew T. & Shultz, Jeffrey W. (2004). "Elastic energy storage in the pedipedal joints of scorpions and sun-spiders (Arachnida, Scorpiones, Solifugae)". Journal of Arachnology . 32 (1): 1–10. doi:10.1636/S02-73. ISSN   0161-8202. S2CID   56461501.
  13. Garwood, Russell J. & Edgecombe, Gregory D. (September 2011). "Early Terrestrial Animals, Evolution, and Uncertainty". Evolution: Education and Outreach. 4 (3): 489–501. doi: 10.1007/s12052-011-0357-y .
  14. 1 2 3 4 5 6 7 Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 596–604. ISBN   978-0-03-056747-6.
  15. 1 2 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.
  16. McLean, Callum J.; Garwood, Russell J.; Brassey, Charlotte A. (2018). "Sexual dimorphism in the Arachnid orders". PeerJ. 6: e5751. doi: 10.7717/peerj.5751 . ISSN   2167-8359. PMC   6225839 . PMID   30416880.
  17. 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". Molecular Biology and Evolution. 27 (11): 2451–2464. doi: 10.1093/molbev/msq130 . PMID   20534705.
  18. 1 2 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. doi:10.1038/nature08742. PMID   20147900. S2CID   4427443.
  19. 1 2 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". Proceedings of the Royal Society of London B: Biological Sciences. 278 (1703): 298–306. doi:10.1098/rspb.2010.0590. PMC   3013382 . PMID   20702459.
  20. 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". Proceedings of the National Academy of Sciences. 108 (38): 15920–15924. Bibcode:2011PNAS..10815920C. doi:10.1073/pnas.1105499108. PMC   3179045 . PMID   21896763.
  21. 1 2 3 4 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". Molecular Biology and Evolution. 31 (11): 2963–2984. doi: 10.1093/molbev/msu235 . PMID   25107551 . Retrieved 2016-03-24.
  22. Legg, David A.; Sutton, Mark D. & Edgecombe, Gregory D. (2013). "Arthropod fossil data increase congruence of morphological and molecular phylogenies". Nature Communications. 4: 2485. Bibcode:2013NatCo...4.2485L. doi: 10.1038/ncomms3485 . PMID   24077329.
  23. 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. doi:10.1038/35093097. PMID   11557979. S2CID   4431635.
  24. 1 2 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". Nature Ecology & Evolution. 2 (4): 614–622. doi:10.1038/s41559-017-0449-3. PMID   29403075. S2CID   4239867.
  25. Garwood, R.J.; Dunlop, J.A.; Knecht, B.J.; Hegna, T.A. (2017). "The phylogeny of fossil whip spiders". BMC Evolutionary Biology. 17 (1): 105. doi:10.1186/s12862-017-0931-1. PMC   5399839 . PMID   28431496.
  26. 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". Proceedings of the Royal Society B: Biological Sciences. 283 (1827): 20160125. doi:10.1098/rspb.2016.0125. PMC   4822468 . PMID   27030415.
  27. Garwood, R.J.; Dunlop, J. (2014). "Three-dimensional reconstruction and the phylogeny of extinct chelicerate orders". PeerJ. 2: e641. doi:10.7717/peerj.641. PMC   4232842 . PMID   25405073.
  28. Shultz, J.W. (2007). "A phylogenetic analysis of the arachnid orders based on morphological characters". Zoological Journal of the Linnean Society. 150 (2): 221–265. doi: 10.1111/j.1096-3642.2007.00284.x .
  29. 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, doi:10.1073/pnas.0809174106, PMC   2634869 , PMID   19104044
  30. Briggs, Helen (5 February 2018). "'Extraordinary' fossil sheds light on origins of spiders". BBC. Retrieved 9 June 2018.
  31. Chapman, Arthur D. (2005). Numbers of living species in Australia and the world (PDF). Department of the Environment and Heritage. ISBN   978-0-642-56850-2.