Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Chelicerata
Classis: Arachnida
Ordo: Thelyphonida
Familiae: Hypoctonidae - Geralinuridae - Thelyphonidae
References
Koch, C. L. 1851. Übersicht des Arachnidensystems 5. C. H. Zeh’sche Buchhandlung, Nürnberg, 104 pp.
Latreille, P. A. 1804. Histoire naturelle, generale et particulière, des Crustacés et des Insectes, Vol. 7. F. Dufart, Paris: 144–305.
Prendini, L. 2011. Thelyphonida Latreille, 1804. P 155 In Zhang, Z.-Q. (ed.) 2011. Animal biodiversity: an outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148: 1–237. Open access. Reference page. . (PDF) Reference page.
Thorell, T. 1882. Descrizione di Alcuni Aracnidi Inferiori dell’ Arcipelago Malese. Annali del Museo Civico di Storia Naturale di Genova, 18: 21–69.
Thorell, T. 1888. Pedipalpi e Scorpioni dell’Arcipelago Malese conservati nel Museo Civico di Storia Naturale di Genova. Annali del Museo Civico di Storia Naturale di Genova, 26: 327–428.
Vernacular names
Deutsch: Geißelskorpione
Ελληνικά: Ουρόπυγα
English: Whip scorpions
español: Uropígidos, vinagrillos
français: Uropyges
magyar: Ostorfarkúak
日本語: サソリモドキ
한국어: 식초전갈,채찍전갈
lietuvių: Telifonai
polski: Biczykoodwłokowce
português: Escorpião-vinagre
svenska: Gisselskorpioner
தமிழ்: சவுக்கை தேள்
ไทย: แมงป่องแส้, แมงป่องหางแส้
Thelyphonida is an arachnid order comprising invertebrates commonly known as whip scorpions or vinegaroons (also spelled vinegarroons and vinegarones). They are often called uropygids in the scientific community based on an alternative name for the order, Uropygi (which may then also include the order Schizomida). The name "whip scorpion" refers to their resemblance to true scorpions and possession of a whiplike tail, and "vinegaroon" refers to their ability when attacked to discharge an offensive, vinegar-smelling liquid, which contains acetic acid.
Taxonomy
Thelyphonus doriae hosei
Carl Linnaeus first described a whip scorpion in 1758, although he did not distinguish it from what are now regarded as different kinds of arachnid, calling it Phalangium caudatum. Phalangium is now used as a name for a genus of harvestmen (Opiliones). In 1802, Pierre André Latreille was the first to use a genus name solely for whip scorpions, namely Thelyphonus.[1][2] Latreille later explained the name as meaning "qui tue", meaning "who kills".[3][note 1] One name for the order, Thelyphonida, is based on Latreille's genus name. It was first used (with the spelling Thelyphonidea) by O. P. Cambridge in 1872.[4]
The name "uropygid" means "tail rump", from Ancient Greek οὐροπύγιον (ouropugion),[5] from οὐρά (oura) "tail" and πυγή (puge) "rump" referring to the whip-like flagellum on the end of the pygidium, a small plate made up of the last three segments of the abdominal exoskeleton.
The classification and scientific name used for whip scorpions varies. Originally, Amblypygi (whip spiders), Thelyphonida and Schizomida (short-tailed whipscorpions) formed a single order of arachnids, Pedipalpi. Pedipalpi was later divided into two orders, Amblypygi and Uropygi (or Uropygida). Schizomida was then split off from Uropygi into a separate order.[6] The remainder has either continued to be called by the same name, Uropygi,[1] possibly distinguished as Uropygi sensu stricto, or called Thelyphonida.[6] Other sources do not accept the split, and continue to treat Schizomida as part of Uropygi.[7] Phylogenetic studies show the three orders to be closely related, and the older order names may now be used as names for clades, as shown below.[6][8] The Schizomida and Thelyphonida likely diverged in the late Carboniferous, somewhere in the tropics of Pangaea.[9]
| Pedipalpi |
|
Description
Whip scorpions range from 25 to 85 mm (1.0 to 3.3 in) in length, with most species having a body no longer than 30 mm (1.2 in); the largest species, of the genus Mastigoproctus, can reach 85 mm (3.3 in).[10] An extinct Mesoproctus from the Lower Cretaceous Crato Formation could be the same size.[11] Because of their legs, claws, and "whip", though, they can appear much larger, and the heaviest specimen weighed was 12.4 grams (0.44 oz).[12]
The opisthosoma consist of 12 segments. The first segment forms a pedicel, and the next eight segments each has dorsal tergits. The last three segments are fused into closed rings that ends with the flagellum, made up of 30-40 units.[13][14]
Like the related orders Schizomida and Amblypygi, the vinegaroons use only six legs for walking, with the first two legs serving as antennae-like sensory organs. All species also have very large scorpion-like pedipalps (pincers) but there is an additional large spine on each palpal tibia. They have one pair of median eyes at the front of the cephalothorax and up till five pairs of lateral eyes on each side of the head, a pattern also found in scorpions.[15][16] Vinegaroons have no venom glands, but they have glands near the rear of their abdomen that can spray a combination of acetic acid and caprylic acid when they are bothered.[10] The acetic acid gives this spray a vinegar-like smell, giving rise to the common name vinegaroon.
Behaviour
Vinegaroons are carnivorous, nocturnal hunters feeding mostly on insects, millipedes, scorpions, and terrestrial isopods,[10] but sometimes on worms and slugs. Mastigoproctus sometimes preys on small vertebrates.[10] The prey is crushed between special teeth on the inside of the trochanters (the second segment of the "legs") of the front appendages. They are valuable in controlling the population of cockroaches and crickets.
Males secrete a spermatophore (a united mass of sperm), which is transferred to the female following courtship behaviour, in which the male holds the ends of the female's first legs in his chelicerae. The spermatophore is deposited on the ground and picked up by the female using her genital area. In some genera, the male then uses his pedipalps to push the spermatophore into her body.[7]
After a few months, the female will dig a large burrow and seal herself inside. Up to 40 eggs are extruded, within a membranous broodsac that preserves moisture and remains attached to the genital operculum and the fifth segment of the mother's ventral opisthosoma. The female refuses to eat and holds her opisthosoma in an upward arch so that the broodsac does not touch the ground for the next few months, as the eggs develop into postembryos. Appendages become visible.[17]
The white young that hatch from the postembryos climb onto their mother's back and attach themselves there with special suckers. After the first molt, when they look like miniature adults but with bright red palps, they leave the burrow. The mother may live up to two more years. The young grow slowly, going through four molts in about four years before reaching adulthood. They live for up to another four years.[10][17]
Distribution and habitat
Whip scorpions are found in tropical and subtropical areas, excluding Europe and Australia. Also, only a single species is known from Africa: Etienneus africanus, probably a Gondwana relict endemic to Senegal, the Gambia and Guinea-Bissau.[18] They usually dig burrows with their pedipalps, to which they transport their prey.[10] They may also burrow under logs, rotting wood, rocks, and other natural debris. They prefer humid, dark places and avoid light. Mastigoproctus giganteus, the giant whip scorpion, is found in more arid areas, including Arizona and New Mexico.[19]
Subtaxa
Main article: List of Thelyphonidae species
As of 2022, the World Uropygi Catalog accepts the following twenty-three genera:[20]
Mastigoproctus giganteus female with eggs
Etienneus Heurtault, 1984
Ginosigma Speijer, 1936
Glyptogluteus Rowland, 1973
Hypoctonus Thorell, 1888
Labochirus Pocock, 1894
Mastigoproctus Pocock, 1894
Mayacentrum Viquez & Armas, 2006
Mimoscorpius Pocock, 1894
Ravilops Víquez & Armas, 2005
Sheylayongium Teruel, 2018
Thelyphonellus Pocock, 1894
Thelyphonoides Krehenwinkel, Curio, Tacud & Haupt, 2009
Thelyphonus Latreille, 1802
Typopeltis Pocock, 1894
Uroproctus Pocock, 1894
Valeriophonus Viquez & Armas, 2005
†Burmathelyphonia Wunderlich, 2015
†Geralinura Scudder, 1884
†Mesoproctus Dunlop, 1998
†Mesothelyphonus Cai & Huang, 2017
†Parageralinura Tetlie & Dunlop, 2008
†Proschizomus Dunlop & Horrocks, 1996
†Prothelyphonus Frič, 1904
Notes
In Greek φόνος, phonos, means "murder", while φονός, with final accent, can be an adjective meaning "murderous", but also a noun meaning "murderess"; Latreille did not account for the element θῆλυς, thelys, meaning "female".
References
Harvey, M.S. (2002). "The neglected cousins: what do we know about the smaller arachnid orders?" (PDF). Journal of Arachnology. 30 (2): 357–372. doi:10.1636/0161-8202(2002)030[0357:TNCWDW]2.0.CO;2. S2CID 59047074. Archived from the original (PDF) on 2012-02-07.
Latreille, Pierre A. (1802). "Genre Thélyphone". Histoire naturelle générale et particulière des Crustacés et des Insectes (in French). Vol. 3. Paris: Dufart. p. 47.
Latreille, Pierre A. (1804). "Genre Thélyphone". Histoire naturelle générale et particulière des Crustacés et des Insectes (in French). Vol. 7. Paris: Dufart. pp. 130–132.
Cambridge, O.P. (1872). "On a new family and genus and two new species of Thelyphonidea". Annals and Magazine of Natural History. 4. 10 (60): 409–413. doi:10.1080/00222937208696729. Retrieved 2016-04-03.
Found in Aristoteles' work: De Anim. Hist., Lib: IV Cap: I.
Garwood, Russell J.; Dunlop, Jason A. (2014). "Three-dimensional reconstruction and the phylogeny of extinct chelicerate orders". PeerJ. 2: e641. doi:10.7717/peerj.641. PMC 4232842. PMID 25405073.
Ruppert, E.E.; Fox, R.S. & Barnes, R.D. (2004). Invertebrate Zoology (7th ed.). Brooks/Cole. pp. 569–570. ISBN 978-0-03-025982-1.
Shultz, Jeffrey 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.
Clouse, Ronald M.; Branstetter, Michael G.; Buenavente, Perry; Crowley, Louise M.; Czekanski‐Moir, Jesse; General, David Emmanuel M.; Giribet, Gonzalo; Harvey, Mark S.; Janies, Daniel A. (2017). "First global molecular phylogeny and biogeographical analysis of two arachnid orders (Schizomida and Uropygi) supports a tropical Pangean origin and mid-Cretaceous diversification". Journal of Biogeography. 44 (11): 2660–2672. doi:10.1111/jbi.13076. ISSN 1365-2699.
Günther Schmidt (1993). Giftige und gefährliche Spinnentiere [Poisonous and dangerous arachnids] (in German). Westarp Wissenschaften. ISBN 978-3-89432-405-6.
Jason A. Dunlop, David M. Martill. The first whipspider (Arachnida: Amblypygi) and three new whipscorpions (Arachnida: Thelyphonida) from the Lower Cretaceous Crato Formation of Brazil. Transactions of the Royal Society of Edinburgh: Earth Sciences, 92, 325–334, 2002.
Glenday, Craig (2013). Guinness World Records 2014. pp. 33. ISBN 978-1-908843-15-9.
Biological Science Fundamentals and Systematics - Volum III
Multicellular Animals: Volume II: The Phylogenetic System of the Metazoa
Ecology and Evolution of the Acari
Invertebrates, by Richard C. Brusca and Gary J. Brusca, page 505
McMonigle, Orin (1 November 2017). Whipscorpions and Whipspiders Culturing Gentle Monsters. Elytra and Antenna. ISBN 978-0980240122.
Jeremy C. Huff & Lorenzo Prendini (2009). "On the African whip scorpion, Etienneus africanus (Hentschel, 1899) (Thelyphonida: Thelyphonidae), with a redescription based on new material from Guinea-Bissau and Senegal". American Museum Novitates (3658): 1–16. doi:10.1206/674.1. hdl:2246/5981. S2CID 59942800.
"Giant whip scorpion Mastigoproctus giganteus giganteus (Lucas, 1835)". Featured Creatures. Institute of Food and Agricultural Sciences, University of Florida. Retrieved 2016-08-01.
"World Uropygi Catalog". World Uropygi Catalog. Natural History Museum Bern. 2022. Retrieved 28 September 2022.
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