Mygalomorphae

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Mygalomorphae
Temporal range: Anisian–Recent
Mouse spider.jpg
Missulena bradleyi, a mouse spider
Scientific classification e
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Order: Araneae
Suborder: Opisthothelae
Infraorder: Mygalomorphae
Pocock, 1892[1]
Subdivisions

The Mygalomorphae, or mygalomorphs, are an infraorder of spiders, and comprise one of three major groups of living spiders with over 3000 species, found on all continents except Antarctica. Many members are known as trapdoor spiders due to them forming trapdoors over their burrows. Other prominent groups include Australian funnel web spiders, and tarantulas, with the latter accounting for around one third of all mygalomorphs.

Description[edit]

This group of spiders comprises mostly heavy-bodied, stout-legged spiders including tarantulas, Australian funnel-web spiders, mouse spiders, and various families of spiders commonly called trapdoor spiders.

Like the "primitive" suborder of spiders Mesothelae, they have two pairs of book lungs, and downward-pointing chelicerae. Because of this, the two groups were once believed to be closely related. Later it was realized that the common ancestors of all spiders had these features (a state known as symplesiomorphy). Following the branching into the suborders of Mesothelae and Opisthothelae, the mygalomorphs retained them, while their fellow Opisthothelae members, the araneomorphs, evolved new "modern" features, including a cribellum and cross-acting fangs.[2] Mesotheles retain the external abdominal segmentation of ancestral arachnids and have at least vestiges of four pairs of spinnerets, whereas mygalomorphs lack abdominal segmentation (like other opistotheles) and have a reduced number of spinnerets, often only two pairs.[3]

Like spiders in general, most species of Mygalomorphae have eight eyes, one pair of principal and three pairs of secondary eyes.

Their chelicerae and fangs are large and powerful and have ample venom glands that lie entirely within their chelicerae. These weapons, combined with their size and strength, make Mygalomorph spiders powerful predators. Many of these spiders are well adapted to killing other large arthropods and will also sometimes kill small mammals, birds, and reptiles. Despite their fearsome appearance and reputation, most mygalomorph spiders are not harmful to humans, with the exception of the Australian funnel-web spiders, especially those of the genus Atrax.[citation needed]

While the world's biggest spiders are mygalomorphs – Theraphosa blondi has a body length of 10 cm (3.9 in) and a leg span of 28 cm (11 in) – some species are less than one millimeter (0.039 in) long. Mygalomorphs are capable of spinning at least slightly adhesive silk, and some build elaborate capture webs that approach a meter in diameter.[2]

Unlike Araneomorphae, which die after about a year, Mygalomorphae can live for up to 25 years, and some do not reach maturity until they are about six years old.[4] Some flies in the family Acroceridae that are endoparasites of mygalomorphs may remain dormant in their book lungs for as long as 20 years before beginning their development and consuming the spider.[citation needed]

One female trapdoor spider, first recorded in a survey in 1974 in Western Australia, is known to have lived for 43 years.[5]

Taxonomy[edit]

The name is derived from the Greek mygalē, meaning "shrew", plus morphē meaning form or shape.[6] An older name for the group is Orthognatha, derived from the orientation of the fangs which point straight down and do not cross each other (as they do in the araneomorphs).

Evolution[edit]

Megarachne servinei was thought to be a giant mygalomorph from the Upper Carboniferous (about 350 million years ago), but was later found to be a eurypterid.[7] The oldest known mygalomorph is Rosamygale grauvogeli , an avicularoid from the Triassic of northeastern France. No mygalomorphs from the Jurassic have yet been found.[8]

The number of families and their relationships have both been undergoing substantial changes since a cladogram showing family relationships was published in 2005,[9] with two significant studies in 2018.[10][11] The division of Mygalomorphae into two superfamilies, Atypoidea and Avicularioidea, has been established in many studies. The Atypoidea retain some vestiges of abdominal segmentation in the form of dorsal tergites; the Avicularioidea lack these. Molecular phylogenetic studies undertaken between 2012 and 2017 have found somewhat different relationships within the Avicularioidea. Some families appear not to be monophyletic and further changes are possible in the future.[3] Rosamygale belongs Avicularioidea, based on the absence of an abdominal scutum and well-separated posterior lateral spinnerets.[12]

Mygalomorphae tend to be highly morphologically conserved, which makes it difficult to find reliable morphological features to use for taxonomy. It has been hypothesized that because Mygalomorphae all tend to be fossorial and live in tubular webs, they are subjected to similar selective pressures, so most species should evolve in similar ways. Additionally, this may also mean that homoplasies are more likely to occur, further complicating taxonomy based on morphology.[13]

Phylogeny[edit]

The relationships of taxa in the Mygalomorphae were restructured based on a comprehensive phylogenetic study by Opatova et al. (2020)[14] The generic composition of the families Ctenizidae, Cyrtaucheniidae, Dipluridae, and Nemesiidae were relimited. Five subfamilies were raised to the rank of family: Anamidae, Euagridae, Ischnothelidae, Pycnothelidae, and Bemmeridae. Three new families were created: Entypesidae, Microhexuridae, and Stasimopidae. Lastly, a new subfamily, Australothelinae, was generated and placed in the family Euagridae.

The preferred cladogram from Optova et al. (2020) is:[14]

Mygalomorphae
Atypoidea

Hexurellidae

Mecicobothriidae

Atypidae

Megahexuridae

Antrodiaetidae

Avicularioidea

Ischnothelidae

Microhexuridae

Hexathelidae

Euagridae

Porrhothelidae

Macrothelidae

Bipectina

Paratropididae

Stasimopidae

Venom Clade

Atracidae

Actinopodidae

Domiothelina

Halonoproctidae

Migidae

Idiopidae

Ctenizidae

Euctenizidae

Crassitarsae
Theraphosoidina

Bemmeridae

Barychelidae

Theraphosidae

"Nemesioidina"

Nemesiidae

Pycnothelidae

Dipluridae

Cyrtaucheniidae

Anamidae

Entypesidae

Microstigmatidae

Families[edit]

Key
Genera 1 ≥2 ≥10 ≥100
Species 1–9 ≥10 ≥100 ≥1000
Mygalomorphae families[a]
Family Genera Species Common name Example
Atypoidea:[3]
Antrodiaetidae 4 37 folding trapdoor spiders Atypoides riversi
Atypidae 3 54 atypical tarantulas or purseweb spiders Sphodros rufipes (red legged purseweb spider)
Hexurellidae 1 4 Hexurella pinea
Mecicobothriidae 1 2 dwarf tarantulas or sheet funnel-web spiders Mecicobothrium thorelli
Megahexuridae 1 1 Megahexura fulva
Avicularioidea:[3]
Actinopodidae 3 115 Missulena bradleyi (Eastern mouse spider)
Anamidae 10 109 Aname atra
Atracidae 3 35 Australian funnel-web spiders Atrax robustus (Sydney funnel-web spider)
Barychelidae 40 282 Brushed trapdoor spiders Sason sundaicum
Bemmeridae 4 47 Spiroctenus personatus
Ctenizidae 2 5 cork-lid trapdoor spiders Cteniza sauvagesi
Cyrtaucheniidae 8 107 wafer trapdoor spiders Fufius lucasae
Dipluridae 7 92 Curtain-web spiders Diplura lineata
Entypesidae 3 37 Entypesa andohahela
Euagridae 13 86 Euagrus formosanus
Euctenizidae 8 77 Aptostichus simus
Halonoproctidae 6 119 Bothriocyrtum californicum (California trapdoor spider)
Hexathelidae 7 45 (Australian) funnel-web spiders Hexathele hochstetteri
Idiopidae 23 414 armored trapdoor spiders Idiosoma nigrum (black rugose trapdoor spider)
Ischnothelidae 5 26 Ischnothele caudata
Macrothelidae 1 40 Macrothele calpeiana (Spanish funnel-web spider)
Microhexuridae 1 2 Microhexura montivaga (Spruce-fir moss spider)
Microstigmatidae 12 41 Envia garciai
Migidae 11 102 tree trapdoor spiders Calathotarsus simoni
Nemesiidae 22 184 Amblyocarenum nuragicus
Paratropididae 4 16 baldlegged spiders Paratropis tuxtlensis
Porrhothelidae 1 5 Porrhothele antipodiana (black tunnelweb spider)
Pycnothelidae 7 83 Stanwellia hoggi
Stasimopidae 1 47 Stasimopus mandelai
Theraphosidae 152 1009 tarantulas Theraphosa blondi (Goliath birdeater)

Historical classification[edit]

In 1802, C. A. Walckenaer separated mygalomorph spiders into a separate genus, Mygale, leaving all other spiders in Aranea

In 1985, Robert Raven published a monograph of the Mygalomorphae in which he proposed an internal classification for the Mygalomorphae, based on morphological features. Opatova et al. (2020) commented "In short, much of today’s classification scheme dates back to Raven (1985)".[14] Raven used various compound ranks, such as "gigapicoorder" and "hyperpicoorder". Ignoring these unusual rank names, his classification can be shown diagrammatically:[16]

Tuberculotae
Mecicobothrioidina

Mecicobothriidae

Microstigmatidae

Orthopalpae
Hexatheloidina

Hexathelidae

Quadrithelina
Dipluroida

Dipluridae

Crassitarsae
Nemesioida

Nemesiidae

Theraphosoidina
Superfamily Theraphosoidea

Theraphosidae

Paratropididae

Superfamily Barycheloidea

Barychelidae

Fornicephalae
Atypoidina

Atypidae

Antrodiaetidae

Rastelloidina
Cyrtaucheniina

Cyrtaucheniidae

Domiothelina
Idiopina

Idiopidae

Ctenizoidina
Superfamily Ctenizoidea

Ctenizidae

Superfamily Migoidea

Migidae

Actinopodidae

Subsequent research, largely based on molecular phylogenetic studies, has not upheld some of Raven's groupings. In particular his primary division between Tuberculotae and Fornicephalae has been replaced by a very different division between Atypoidea (expanded from Raven's Atypoidina) and Avicularioidea, which has no counterpart in his system. As another example, the families Mecicobothriidae and Microstigmatidae, which Raven placed in Mecicobothrioidina (a "gigapicoorder"), are now placed very far apart in the Atypoidea and Avicularioidea respectively. Other groups, such as Crassitarsae and Domiothelina, are more recognizable, allowing for some changes in family circumscriptions.[14]

Distribution[edit]

Most members of this infraorder occur in the tropics and subtropics, but their range can extend farther north, e.g. into the southern and western regions of the United States. Only a few occur in Europe: 12 species from the families Atypidae, Nemesiidae, Ctenizidae, Macrothelidae, Theraphosidae, and Cyrtaucheniidae.

Despite their limited range, currently, it is suggested that the Mygalomorphae were distributed worldwide before the breakup of Pangaea.[8]

Notes[edit]

  1. ^ Unless otherwise shown, currently accepted families and counts based on the World Spider Catalog version 22.0 as of 21 April 2021.[15] In the World Spider Catalog, "species" counts include subspecies. Assignment to sub- and infraorders based on Coddington (2005, p. 20) (when given there).

References[edit]

  1. ^ Dunlop, Jason A. & Penney, David (2011). "Order Araneae Clerck, 1757" (PDF). In Zhang, Z.-Q. (ed.). Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa. Auckland, New Zealand: Magnolia Press. ISBN 978-1-86977-850-7. Retrieved 2015-10-31.
  2. ^ a b Coddington, Jonathan A. & Levi, Herbert W. (1991). "Systematics and evolution of spiders (Araneae)". Annual Review of Ecology and Systematics. 22: 565–592. doi:10.1146/annurev.es.22.110191.003025. JSTOR 2097274. S2CID 55647804.
  3. ^ a b c d Wheeler, Ward C.; Coddington, Jonathan A.; Crowley, Louise M.; Dimitrov, Dimitar; Goloboff, Pablo A.; Griswold, Charles E.; et al. (2016). "The spider tree of life: Phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling". Cladistics. 33 (6): 574–616. doi:10.1111/cla.12182. S2CID 35535038.
  4. ^ "About Spiders". CSIRO. Archived from the original on 2015-09-24. Retrieved 2017-02-18.
  5. ^ "World's oldest spider dies aged 43 in Western Australia". ABC News. Australian Broadcasting Corporation. 28 April 2018. Archived from the original on 2018-04-29. Retrieved 2018-04-29.
  6. ^ "mygalomorph". Oxford Dictionaries. Archived from the original on 2016-05-21. Retrieved 2016-02-02.
  7. ^ Selden, P.A.; Corronca, J.A. & Hünicken, M.A. (2005). "The true identity of the supposed giant fossil spider Megarachne". Biology Letters. 1 (1): 44–48. doi:10.1098/rsbl.2004.0272. PMC 1629066. PMID 17148124.
  8. ^ a b Selden, P.A.; da Costa Casado, F. & Vianna Mesquita, M. (2005). "Mygalomorph spiders (Araneae: Dipluridae) from the Lower Cretaceous Crato Lagerstätte, Araripe Basin, north-east Brazil". Palaeontology. 49 (4): 817–826. doi:10.1111/j.1475-4983.2006.00561.x. hdl:1808/8355.
  9. ^ Coddington, Jonathan A. (2005). "Phylogeny and classification of spiders" (PDF). In Ubick, D.; Paquin, P.; Cushing, P.E. & Roth, V. (eds.). Spiders of North America: an identification manual. American Arachnological Society. pp. 18–24. Retrieved 2015-09-24.
  10. ^ Hedin, Marshal; Derkarabetian, Shahan; Ramírez, Martín J.; Vink, Cor & Bond, Jason E. (2018). "Phylogenomic reclassification of the world's most venomous spiders (Mygalomorphae, Atracinae), with implications for venom evolution". Scientific Reports. 8 (1): 1636. doi:10.1038/s41598-018-19946-2. PMC 5785998. PMID 29374214.
  11. ^ Godwin, Rebecca L.; Opatova, Vera; Garrison, Nicole L.; Hamilton, Chris A. & Bond, Jason E. (2018). "Phylogeny of a cosmopolitan family of morphologically conserved trapdoor spiders (Mygalomorphae, Ctenizidae) using Anchored Hybrid Enrichment, with a description of the family, Halonoproctidae (Pocock, 1901)". Molecular Phylogenetics and Evolution. 126: 303–313. doi:10.1016/j.ympev.2018.04.008. ISSN 1055-7903. PMID 29656103.
  12. ^ Magalhaes, Ivan L.F.; Azevedo, Guilherme H.F.; Michalik, Peter; Ramírez, Martín J. (2020). "The fossil record of spiders revisited: Implications for calibrating trees and evidence for a major faunal turnover since the Mesozoic". Biological Reviews. 95 (1): 184–217. doi:10.1111/brv.12559. ISSN 1464-7931. PMID 31713947. S2CID 207937170.
  13. ^ Bond, Jason E.; Hedin, Marshal (1 October 2006). "A total evidence assessment of the phylogeny of North American euctenizine trapdoor spiders (Araneae, Mygalomorphae, Cyrtaucheniidae) using Bayesian inference". Molecular Phylogenetics and Evolution. 41 (1): 70–85. doi:10.1016/j.ympev.2006.04.026. ISSN 1055-7903. PMID 16857391.
  14. ^ a b c d Opatova, Vera; Hamilton, Chris A.; Hedin, Marshal; Montes De Oca, Lauren; Král, Jiři; Bond, Jason E. (16 October 2019). "Phylogenetic Systematics and Evolution of the Spider Infraorder Mygalomorphae Using Genomic Scale Data". Systematic Biology. 69 (4): 671–707. doi:10.1093/sysbio/syz064. PMID 31841157.
  15. ^ "Currently valid spider genera and species". World Spider Catalog. Natural History Museum Bern. Retrieved 2021-04-21.
  16. ^ Raven, Robert J. (1985). "The spider infraorder Mygalomorphae (Araneae) : cladistics and systematics". Bulletin of the American Museum of Natural History. 182 (1): 1–180.

Bibliography[edit]

  • Raven, R.J. (1985). "The spider infraorder Mygalomorphae: Cladistics and systematics". Bulletin of the American Museum of Natural History. 182: 1–180.

External links[edit]