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Thelyphonida

February 18, 2023

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.

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

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

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

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

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".

References

  1. ^ a b Harvey, M.S. (2002). (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. ^ a b c 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. ^ a b 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. ^ a b c d e f 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. ^ a b 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.

External links

 
Wikimedia Commons has media related to Thelyphonida.
  • Video of vinegaroon mating behavior

February 18, 2023
thelyphonida, arachnid, order, comprising, invertebrates, commonly, known, whip, scorpions, vinegaroons, also, spelled, vinegarroons, vinegarones, they, often, called, uropygids, scientific, community, based, alternative, name, order, uropygi, which, then, als. 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 Whip scorpions vinegaroonsTemporal range Carboniferous present PreꞒ Ꞓ O S D C P T J K Pg NScientific classificationKingdom AnimaliaPhylum ArthropodaSubphylum ChelicerataClass ArachnidaClade TetrapulmonataOrder ThelyphonidaO P Cambridge 1872FamiliesThelyphonidaeDiversityc 23 genera gt 100 speciesWhip scorpion in Florida Contents 1 Taxonomy 2 Description 3 Behaviour 4 Distribution and habitat 5 Subtaxa 6 Notes 7 References 8 External linksTaxonomy Edit 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 Andre 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 oὐropygion ouropugion 5 from oὐra oura tail and pygh 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 EditWhip 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 EditVinegaroons 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 EditWhip 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 EditMain 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 amp Armas 2006 Mimoscorpius Pocock 1894 Ravilops Viquez amp Armas 2005 Sheylayongium Teruel 2018 Thelyphonellus Pocock 1894 Thelyphonoides Krehenwinkel Curio Tacud amp Haupt 2009 Thelyphonus Latreille 1802 Typopeltis Pocock 1894 Uroproctus Pocock 1894 Valeriophonus Viquez amp Armas 2005 Burmathelyphonia Wunderlich 2015 Geralinura Scudder 1884 Mesoproctus Dunlop 1998 Mesothelyphonus Cai amp Huang 2017 Parageralinura Tetlie amp Dunlop 2008 Proschizomus Dunlop amp Horrocks 1996 Prothelyphonus Fric 1904Notes Edit In Greek fonos phonos means murder while fonos with final accent can be an adjective meaning murderous but also a noun meaning murderess Latreille did not account for the element 8ῆlys thelys meaning female References Edit a b 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 Thelyphone Histoire naturelle generale et particuliere des Crustaces et des Insectes in French Vol 3 Paris Dufart p 47 Latreille Pierre A 1804 Genre Thelyphone Histoire naturelle generale et particuliere des Crustaces 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 a b c 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 a b Ruppert E E Fox R S amp 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 a b c d e f Gunther Schmidt 1993 Giftige und gefahrliche 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 a b McMonigle Orin 1 November 2017 Whipscorpions and Whipspiders Culturing Gentle Monsters Elytra and Antenna ISBN 978 0980240122 Jeremy C Huff amp 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 External links Edit Wikimedia Commons has media related to Thelyphonida Arthropods portalVideo of vinegaroon mating behavior Retrieved from https en wikipedia org w index php title Thelyphonida amp oldid 1130164854, wikipedia, wiki, book, 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