Thelyphonida

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Whip scorpions, vinegaroons
Temporal range: Carboniferous–present
Whipscorpion.jpg
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Order: Thelyphonida
O. P-Cambridge, 1872
Families
Diversity
c. 23 genera, > 100 species
Whip scorpion in Florida Whip scorpion.jpg
Whip scorpion in Florida

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.

Contents

Taxonomy

Thelyphonus doriae hosei WhipscorpLyd.png
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]

Liuzhou Thelyphonida.jpg
Liuzhou Whip Scorpion.jpg

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

Amblypygi (whip spiders)

Uropygi sensu lato

Thelyphonida (Uropygi sensu stricto, whip scorpions)

Schizomida (short-tailed whip scorpions)

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

As of 2022, the World Uropygi Catalog accepts the following twenty-three genera: [20]

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

Notes

  1. ↑ 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

Arachnida is a class of joint-legged invertebrate animals (arthropods), in 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 comprising whip spiders and tailless whip scorpions

Amblypygi is an ancient order of arachnid chelicerate arthropods also known as African cave-dwelling spiders, 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. 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.

<span class="mw-page-title-main">Schizomida</span> Order of shorttailed whipscorpions

Schizomida is an order of arachnids, generally less than 5 millimetres (0.20 in) in length.

<span class="mw-page-title-main">Pedipalp</span> 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.

<span class="mw-page-title-main">Solifugae</span> Order of spider-like animals

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 147 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 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 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.

<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.

<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.

<i>Chelypus</i> Genus of spider-like animals

Chelypus ('clawfoot') is a genus of slow-moving, burrowing sunspiders confined to the deserts and arid regions of Southern Africa. There are 6 known species.

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

Thelyphonidae is the sole family in the Thelyphonida order of arachnids.

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

Paraphrynus is a genus of whip spiders, also known as tailless whip scorpions, of the family Phrynidae. It is distributed from the southwestern United States to Central America, including several Caribbean islands. Most species are endemic to Mexico.

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

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>Paracharon</i> Genus of whip scorpions

Paracharon is a genus of tailless whip scorpion. A single species, Paracharon caecus has been described. It is endemic to Guinea-Bissau. An undescribed species is known from Colombia. It is the most basal known living member of Amblypygi, and the only living member of the Palaeoamblypygi, which also includes the genus Weygoldtina from the Carboniferous of Euramerica and Paracharonopsis from Eocene aged Indian Cambay amber. Both living species are troglobites, having no eyes, with P. caecus living in termite nests, while the undescribed Colombian species was found living in a cave.

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

Paracharontidae is an arachnid family within the order of tailless whip scorpions. Paracharontidae and Weygoldtinidae form the suborder Paleoamblypygi, the putative sister group to the remaining Amblypygi; this suborder is primarily known from Late Carboniferous and Eocene deposits, but also comprises two extant species: Paracharon caecus Hansen, 1921 from Guinea-Bissau, and an undescribed species from Colombia.

References

  1. 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 2012-02-07.
  2. ↑ Latreille, Pierre A. (1802). "Genre Thélyphone". Histoire naturelle générale et particulière des Crustacés et des Insectes (in French). Vol. 3. Paris: Dufart. p. 47.
  3. ↑ 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.
  4. ↑ 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.
  5. ↑ Found in Aristoteles' work: De Anim. Hist., Lib: IV Cap: I.
  6. 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.
  7. 1 2 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.
  8. ↑ 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 .
  9. ↑ 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.
  10. 1 2 3 4 5 6 Günther Schmidt (1993). Giftige und gefährliche Spinnentiere[Poisonous and dangerous arachnids] (in German). Westarp Wissenschaften. ISBN   978-3-89432-405-6.
  11. ↑ 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.
  12. ↑ Glenday, Craig (2013). Guinness World Records 2014. pp.  33. ISBN   978-1-908843-15-9.
  13. ↑ Biological Science Fundamentals and Systematics - Volum III
  14. ↑ Multicellular Animals: Volume II: The Phylogenetic System of the Metazoa
  15. ↑ Ecology and Evolution of the Acari
  16. ↑ Invertebrates, by Richard C. Brusca and Gary J. Brusca, page 505
  17. 1 2 McMonigle, Orin (1 November 2017). Whipscorpions and Whipspiders Culturing Gentle Monsters. Elytra and Antenna. ISBN   978-0980240122.
  18. ↑ 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.
  19. ↑ "Giant whip scorpion Mastigoproctus giganteus giganteus (Lucas, 1835)". Featured Creatures. Institute of Food and Agricultural Sciences, University of Florida. Retrieved 2016-08-01.
  20. ↑ "World Uropygi Catalog". World Uropygi Catalog. Natural History Museum Bern. 2022. Retrieved 28 September 2022.
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