Uropygi

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Whip scorpions, vinegaroons
Temporal range: Carboniferous–present
Typopeltis crucifer at Fuxing, Taoyuan 2022-06-12 (cropped).jpg
Typopeltis crucifer
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Clade: Tetrapulmonata
Order: Uropygi
Thorell, 1883 [1] [2]
Families
Diversity
c. 23 genera, over 100 species
Uropygi Distribution Map.png
Distribution map of Uropygi
Synonyms [1] [2]

Thelyphonida Latreille, 1804 (as Thélyphone)

Uropygi is an arachnid order comprising invertebrates commonly known as whip scorpions or vinegaroons (also spelled vinegarroons and vinegarones). They are often called uropygids. 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. The order may also be called Thelyphonida. Both names, Uropygi and Thelyphonida, may be used either in a narrow sense for the order of whip scorpions, or in a broad sense which includes the order Schizomida. [a]

Contents

Taxonomy

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 . [3] [4] Latreille later explained the name as meaning "qui tue", meaning "who kills". [5] [b] One name for the order, Thelyphonida, is based on Latreille's genus name. It was first used, as the French Thélyphone, by Latreille in 1804, [1] and later by Octavius Pickard-Cambridge in 1872 (with the spelling Thelyphonidea). [6]

The alternative name, Uropygi, was first used by Tamerlan Thorell in 1883. [1] It means "tail rump", from Ancient Greek οὐροπύγιον (ouropugion), [7] 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), Uropygi 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. [8] The remainder has either continued to be called by the same name, Uropygi, [3] possibly distinguished as Uropygi sensu stricto , or called Thelyphonida. [8] When the name Uropygi is used for the whip scorpions, the clade containing Uropygi and Schizomida may be called Thelyphonida, [9] or Thelyphonida s.l. Conversely, when the name Thelyphonida is used for the whip scorpions alone, the parent clade may be called Uropygi, [10] or Uropygi sensu lato . The table below summarizes the two usages. When the qualifications s.l. and s.s. are omitted, the names Uropygi and Thelyphonida are ambiguous.

Alternative nomenclature
English namesSystem 1System 2Recent families
Thelyphonida s.l. (if used)Uropygi s.l. (if used)
whip scorpions   Uropygi s.s.   Thelyphonida s.s. Thelyphonidae
short-tailed whip scorpions   Schizomida   Schizomida Hubbardiidae, Protoschizomidae

Phylogenetic studies show the three groups, Amblypygi, Uropygi s.s. and Schizomida, to be closely related. [8] [11] The Uropygi s.s. and Schizomida likely diverged in the late Carboniferous, somewhere in the tropics of Pangaea. [12]

  Pedipalpi  
  Amblypygi  

whip spiders Flickr - ggallice - Tailless whip-scorpion, La Muerta.png

  Thelyphonida s.l. 
Uropygi s.s.

whip scorpions Whip Scorpion body (9672115742) (white background).png

  (Thelyphonida s.s.)  
 
  Schizomida  

short-tailed whip scorpions Brignolizomus woodwardi 175486060.jpg

(Uropygi s.l.)

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). [13] An extinct Mesoproctus from the Lower Cretaceous Crato Formation could be the same size. [14] 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). [15]

The opisthosoma consists of 12 segments. The first segment forms a pedicel, and each of the next eight segments has dorsal tergites. The last three segments are fused into closed rings that ends with the flagellum, made up of 30-40 units. [16] [17]

Like the related orders Schizomida and Amblypygi, whip scorpions 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. [18] [19] Whip scorpions 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. [13] The acetic acid gives this spray a vinegar-like smell, giving rise to the common name vinegaroon.

Behaviour

Mastigoproctus giganteus, a predator of millipedes Vinegaroon - Mastigoproctus giganteus with Slate Millipede.jpg
Mastigoproctus giganteus , a predator of millipedes

Whip scorpions are carnivorous, nocturnal hunters feeding mostly on insects, millipedes, scorpions, and terrestrial isopods, [13] but sometimes on worms and slugs. Mastigoproctus sometimes preys on small vertebrates. [13] 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. [20]

Mastigoproctus giganteus female with eggs Mastigoproctus giganteus 0008 L.D.jpg
Mastigoproctus giganteus female with eggs

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. [21]

The white young that hatch from the postembryos climb onto their mother's back and attach themselves there with special suckers. After the first moult, 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 moults in about four years before reaching adulthood. They live for up to another four years. [13] [21]

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. [22] They usually dig burrows with their pedipalps, to which they transport their prey. [13] 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. [23]

Subtaxa

As of 2023, the World Uropygi Catalog accepted the following 16 extant genera, all placed in the family Thelyphonidae: [24]

In addition, seven extinct genera were accepted, two within the family Thelyphonidae: [24]

and five unplaced as to family: [24]

Notes

  1. ↑ For clarity, sensu stricto or s.s. may be added to specify the narrow sense, and sensu lato or s.l. added to specify the broad sense. When these additions are omitted, the names Uropygi and Thelyphonida are ambiguous.
  2. ↑ 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".

Related Research Articles

<span class="mw-page-title-main">Arachnid</span> Class of arthropods

Arachnids are arthropods in the class Arachnida of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaroons.

<span class="mw-page-title-main">Amblypygi</span> Order of arachnids

Amblypygi is an order of arachnids also known as whip spiders or tailless whip scorpions, not to be confused with whip scorpions or vinegaroons that belong to the related order Thelyphonida. The name "amblypygid" means "blunt tail", a reference to a lack of the flagellum that is otherwise seen in whip scorpions. Amblypygids possess no silk glands or venom. They rarely bite if threatened but can grab fingers with their pedipalps, resulting in thorn-like puncture injuries.

<span class="mw-page-title-main">Schizomida</span> Order of short-tailed whip-scorpions

Schizomida, also known as sprickets or short-tailed whip-scorpions, is an order of arachnids, generally less than 5 millimetres (0.20 in) in length. The order is not yet widely studied. E. O. Wilson has identified schizomids as among the "groups of organisms that desperately need experts to work on them."

<i>Plesiosiro</i> Extinct genus of arachnids

Plesiosiro is an extinct arachnid genus known exclusively from 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.

<span class="mw-page-title-main">Trigonotarbida</span> Extinct order of arachnids

The order Trigonotarbida is a group of extinct arachnids whose fossil record extends from the late Silurian to the early Permian. These animals are known from several localities in Europe and North America, as well as a single record from Argentina. Trigonotarbids can be envisaged as spider-like arachnids, but without silk-producing spinnerets. They ranged in size from a few millimetres to a few centimetres in body length and had segmented abdomens (opisthosoma), with the dorsal exoskeleton (tergites) across the backs of the animals' abdomens, which were characteristically divided into three or five separate plates. Probably living as predators on other arthropods, some later trigonotarbid species were quite heavily armoured and protected themselves with spines and tubercles. About seventy species are currently known, with most fossils originating from the Carboniferous coal measures.

<span class="mw-page-title-main">Tetrapulmonata</span> Clade of arachnids

Tetrapulmonata is a non-ranked supra-ordinal clade of arachnids. It is composed of the extant orders Uropygi, 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.

<i>Mastigoproctus</i> Genus of whip scorpions

Mastigoproctus is a genus of whip scorpions. Native to the tropical forest regions of northern South America, these whip scorpions can reach a length of up to 9 centimetres (3.5 in) and can weigh over 30 grams (1.1 oz). Despite popular belief, they are not venomous as, like all other whip scorpions, they do not possess venom glands.

<span class="mw-page-title-main">Charinidae</span> Family of whip scorpions

Charinidae is an arachnid family within the order of tailless whip scorpions.

<i>Mastigoproctus colombianus</i> Species of whip scorpion

Mastigoproctus colombianus is a whip scorpion species found in Colombia, South America, near west deserts areas.

<i>Mastigoproctus giganteus</i> Species of whip scorpion

Mastigoproctus giganteus, the giant whip scorpion, also called the giant vinegaroon or grampus, is a species of whip scorpions in the family Thelyphonidae. Its native range is from the Southern United States to Mexico.

<i>Phrynus</i> Genus of whip scorpions

Phrynus is a genus of whip spiders found in tropical and subtropical regions, mostly in the new world.

<span class="mw-page-title-main">Phrynidae</span> Family of whipspiders

Phrynidae is a family of amblypygid arachnida arthropods also known as whip spiders and tailless whip scorpions. Phrynidae species are found in tropical and subtropical regions in North and South America. Some species are subterranean; all are nocturnal. At least some species of Phrynidae hold territories that they defend from other individuals.

Phrynichus is a genus of tailless whipscorpions in the family Phrynichidae. There are about 16 described species in Phrynichus.

<i>Thelyphonus</i> Genus of whip scorpions

Thelyphonus is the type genus of whip scorpions or 'vinegaroons' in the subfamily Thelyphoninae, with species found in Southeast Asia.

<i>Paracharon</i> Genus of amblypygi

Paracharon is a genus of tailless whip scorpion. A single species, Paracharon caecus has been described. It is endemic to Guinea-Bissau in West Africa It is one of two living genera of the family Paracharontidae, alongside the South American Jorottui. It is a troglobite having no eyes, with P. caecus found living in termite nests.

<span class="mw-page-title-main">Paracharontidae</span> Family of whip scorpions

Paracharontidae is an arachnid family within the order Amblypygi. Paracharontidae and the extinct Weygoldtinidae from the Carboniferous form the suborder Paleoamblypygi, the sister group to the remaining Amblypygi. The family contains two genera: Paracharon, containing the single species Paracharon caecus Hansen, 1921 from Guinea-Bissau in West Africa, and Jorottui with the single species Jorottui ipuanai from Colombia in northern South America. Paracharonopsis from the Eocene (Ypresian) aged Cambay amber of India was initially assigned to this family but this was later questioned and it has since been reassigned to Euamblypygi. Both living species are troglobites, having no eyes, with P. caecus living in termite nests, while J. ipuanai inhabits caves.

<i>Geralinura</i> Extinct genus in the family Thelyphonidae

Geralinura is an extinct genus in the family Thelyphonidae, commonly known as whip scorpions. Fossil evidence found in the midwestern United States and the United Kingdom indicates that this genus lived during the mid-to-late Carboniferous period. Unlike the subchelate pedipalps of modern whip scorpions, the pedipalps of Geralinura are spiked and non-chelate, suggesting that the modern trait evolved at a later point in time.

<i>Mastigoproctus tohono</i> Species of whip scorpion

Mastigoproctus tohono, also known as the Tohono whipscorpion or Tohono vinegaroon, is a species of whip scorpions in the family Uropygi. Its native range is from northern Sonora in Mexico to southern Arizona and western New Mexico, with most sightings occurring in the Chiricahua and Huachuca Mountains of Cochise County, Arizona.

References

  1. 1 2 3 4 Harvey, Mark S. (2003). "Uropygi". Catalogue of the Smaller Arachnid Orders of the World: Amblypygi, Uropygi, Schizomida, Palpigradi, Ricinulei and Solifugae. Canberra, AU: CSIRO Publishing. doi:10.1071/9780643090071. ISBN   978-0-643-09007-1.
  2. 1 2 "Uropygi (Thorell, 1883)". Western Australian Museum . Field Guides & Catalogues. Western Australian Museum, Government of Western Australia . Retrieved 10 March 2023.
  3. 1 2 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 7 February 2012.
  4. ↑ Latreille, Pierre A. (1802). "Genre Thélyphone". Histoire naturelle générale et particulière des Crustacés et des Insectes (in French). Vol. 3. Paris, FR: Dufart. p. 47.
  5. ↑ Latreille, Pierre A. (1804). "Genre Thélyphone". Histoire naturelle générale et particulière des Crustacés et des Insectes (in French). Vol. 7. Paris, FR: Dufart. pp. 130–132.
  6. ↑ 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 3 April 2016.
  7. ↑ Aristotele (28 November 2023). De Anim. Hist. Lib: IV Cap: I.
  8. 1 2 3 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.
  9. ↑ "Order Thelyphonida (Latreille, 1804) (whip scorpion)". Paleobiology Database . Retrieved 10 March 2023.
  10. ↑ "order Uropygi (Thorell, 1883)". BioLib. Retrieved 10 March 2023.
  11. ↑ 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 .
  12. ↑ Clouse, Ronald M.; Branstetter, Michael G.; Buenavente, Perry; Crowley, Louise M.; Czekanski‐Moir, Jesse; General, David Emmanuel M.; et al. (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.
  13. 1 2 3 4 5 6 Schmidt, Günther (1993). Giftige und gefährliche Spinnentiere[Poisonous and dangerous arachnids] (in German). Westarp Wissenschaften. ISBN   978-3-89432-405-6.
  14. ↑ Dunlop, Jason A.; Martill, David M. (2002). "The first whipspider (Arachnida: Amblypygi) and three new whipscorpions (Arachnida: Thelyphonida) from the Lower Cretaceous Crato Formation of Brazil" (PDF). Transactions of the Royal Society of Edinburgh: Earth Sciences. 92 (3): 325–334. doi:10.1017/S0263593300000262. S2CID   140573577.
  15. ↑ Glenday, Craig (2013). Guinness World Records 2014. Guinness World Records Limited. pp.  33. ISBN   978-1-908843-15-9.
  16. ↑ Minelli, Alessandro; Contrafatto, Giancarlo, eds. (10 November 2009). Biological Science Fundamentals and Systematics. Encyclopedia of Life Support Systems. Vol. III. Oxford, UK: EOLSS Publications. p. 225. ISBN   978-1-84826-990-3 – via Google Books.
  17. ↑ Ax, Peter (2000) [1999 original German ed.]. Multicellular Animals: The phylogenetic system of the metazoa. Vol. II: The Phylogenetic System of the Metazoa. Translated by Kiney, S. (English ed.). Springer. p. 103. doi:10.1007/978-3-662-10396-8. ISBN   978-3-642-08681-6. S2CID   28516278 – via Google Books.
  18. ↑ Weygoldt, Peter (1–5 July 1996). "Evolution and systematics of the Chelicerata". In Bruin, J. Leo; van der Geest, P.S.; Sabelis, M.W. (eds.). Ecology and Evolution of the Acari. 3rd Symposium of the European Association of Acarologists. Amsterdam, NL (published 1999). pp. 1–5, esp. p 5. doi:10.1007/978-94-017-1343-6. ISBN   978-90-481-5200-1 – via Google Books.
  19. ↑ Brusca, Richard C.; Brusca, Gary J. Invertebrates (PDF). juristas.com.br (Report). p. 505.
  20. ↑ 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.
  21. 1 2 McMonigle, Orin (November 2017). Whipscorpions and Whipspiders: Culturing gentle monsters. Elytra and Antenna. ISBN   978-098024012-2.
  22. ↑ Huff, Jeremy C.; Prendini, Lorenzo (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.
  23. ↑ "Giant whip scorpion Mastigoproctus giganteus giganteus(Lucas, 1835)". Featured Creatures. Institute of Food and Agricultural Sciences, Department of Entymology. University of Florida . Retrieved 1 August 2016.
  24. 1 2 3 "World Uropygi Catalog". Bern, CH: Natural History Museum. 2024. Retrieved 9 December 2024.
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