Thelyphonida

Thelyphonida is an arachnid order comprising invertebrates

commonly known as whip scorpions or vinegaroons (also spelled Whip scorpions, vinegaroons
vinegarroons and vinegarones). They are often called uropygids Temporal range:
in the scientific community based on an alternative name for the Carboniferous–present
order, Uropygi (which may then also include the order
Schizomida). The name "whip scorpion" refers to their resemblance PreꞒ Ꞓ O S D C P T J K PgN
to true scorpions and possession of a whiplike tail. "Vinegaroon" is
based on their ability when attacked to discharge an offensive liquid
which contains acetic acid, producing a vinegar-like smell.

Contents
Taxonomy
Scientific classification
Description
Kingdom: Animalia
Behaviour
Phylum: Arthropoda
Habitat
Subtaxa Subphylum: Chelicerata
Notes Class: Arachnida
References Order: Thelyphonida
External links O. P-Cambridge, 1872

Families

Taxonomy Thelyphonidae
Carl Linnaeus first described a whip scorpion in 1758, although he Diversity
did not distinguish it from what are now regarded as different kinds
c. 15 genera, > 100 species
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 Whip scorpion in Florida
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]

Amblypygi (whip spiders)

Thelyphonus doriae hosei

Thelyphonida (Uropygi sensu stricto,

Pedipalpi whip scorpions)
Uropygi sensu lato

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] 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).[11]

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 eyes at the front of
the cephalothorax and three on each side of the head, a pattern also found in scorpions.[10] 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.[12]

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, they look like miniature adults but with bright red palps and 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][12]

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.[13] 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.[14]

Subtaxa
As of 2006, over 100 species of vinegaroons have been described worldwide. Subtaxa of vinegaroons
currently include only one extant family and a doubtful extinct family:

Hypoctoninae Pocock, 1899

Etienneus Heurtault, 1984
Hypoctonus Thorell, 1888
Labochirus Pocock, 1894
Thelyphonellus Pocock, 1894

Mastigoproctinae Speijer, 1933

Mastigoproctus Pocock, 1894
Mimoscorpius Pocock, 1894 Mastigoproctus giganteus female
with eggs
Myacentrum Viquez and de Armas, 2006
Uroproctus Pocock, 1894
Valeriophonus Viquez and de Armas, 2005

Thelyphoninae Lucas, 1835

Abaliella Strand, 1928
Chajnus Speijer, 1936
Ginosigma Speijer, 1936
Glyptogluteus Rowland, 1973
Minbosius Speijer, 1933
Tetrabalius Thorell, 1888
Thelyphonus Latreille, 1802

Typopeltinae Rowland & Cooke, 1973

 Typopeltis Pocock, 1894

incertae sedis
† Geralinura Scudder, 1884
† Mesoproctus Dunlop, 1998
† Proschizomus Dunlop & Horrocks, 1996

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. Harvey, M.S. (2002). "The neglected cousins: what do we know about the smaller arachnid
orders?" (https://web.archive.org/web/20120207165733/http://www.americanarachnology.org/J
oA_Congress/JoA_v30_n2/arac-30-02-357.pdf) (PDF). Journal of Arachnology. 30 (2): 357–
372. doi:10.1636/0161-8202(2002)030[0357:TNCWDW]2.0.CO;2 (https://doi.org/10.1636%2F0
161-8202%282002%29030%5B0357%3ATNCWDW%5D2.0.CO%3B2). Archived from the
original (http://www.americanarachnology.org/JoA_Congress/JoA_v30_n2/arac-30-02-357.pdf)
(PDF) on 2012-02-07.
2. Latreille, Pierre A. (1802). "Genre Thélyphone" (https://www.biodiversitylibrary.org/page/24884
903). Histoire naturelle générale et particulière des Crustacés et des Insectes (in French). 3.
Paris: Dufart. p. 47.
3. Latreille, Pierre A. (1804). "Genre Thélyphone" (https://www.biodiversitylibrary.org/page/24883
087). Histoire naturelle générale et particulière des Crustacés et des Insectes (in French). 7.
Paris: Dufart. pp. 130–132.
4. Cambridge, O.P. (1872). "On a new family and genus and two new species of Thelyphonidea"
(https://www.biodiversitylibrary.org/page/25172652). Annals and Magazine of Natural History.
4. 10 (60): 409–413. doi:10.1080/00222937208696729 (https://doi.org/10.1080%2F002229372
08696729). Retrieved 2016-04-03.
5. Found in Aristoteles' work: De Anim. Hist., Lib: IV Cap: I (https://books.google.com/books?id=s
UfySpLBb2wC&pg=PA109-IA3&lpg=PA109-IA3).
6. Garwood, Russell J.; Dunlop, Jason A. (2014). "Three-dimensional reconstruction and the
phylogeny of extinct chelicerate orders" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC423284
2). PeerJ. 2: e641. doi:10.7717/peerj.641 (https://doi.org/10.7717%2Fpeerj.641). PMC 4232842
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4232842). PMID 25405073 (https://pubmed.ncb
i.nlm.nih.gov/25405073).
7. Ruppert, E.E.; Fox, R.S. & Barnes, R.D. (2004). Invertebrate Zoology (https://archive.org/detail
s/isbn_9780030259821/page/569) (7th ed.). Brooks/Cole. pp. 569–570 (https://archive.org/deta
ils/isbn_9780030259821/page/569). ISBN 978-0-03-025982-1. Cite uses deprecated
parameter |lastauthoramp= (help)
8. Shultz, Jeffrey W. (2007). "A phylogenetic analysis of the arachnid orders based on
morphological characters" (https://doi.org/10.1111%2Fj.1096-3642.2007.00284.x). Zoological
Journal of the Linnean Society. 150 (2): 221–265. doi:10.1111/j.1096-3642.2007.00284.x (http
s://doi.org/10.1111%2Fj.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" (https://doi.org/10.1111%2Fjbi.13076). Journal of Biogeography. 44 (11): 2660–
2672. doi:10.1111/jbi.13076 (https://doi.org/10.1111%2Fjbi.13076). ISSN 1365-2699 (https://w
ww.worldcat.org/issn/1365-2699).
10. Günther Schmidt (1993). Giftige und gefährliche Spinnentiere [Poisonous and dangerous
arachnids] (in German). Westarp Wissenschaften. ISBN 978-3-89432-405-6.
11. Glenday, Craig (2013). Guinness World Records 2014 (https://archive.org/details/guinnessworl
drec0000unse_r3e7/page/33). pp. 33 (https://archive.org/details/guinnessworldrec0000unse_r3
e7/page/33). ISBN 978-1-908843-15-9.
12. McMonigle, Orin (1 November 2017). Whipscorpions and Whipspiders Culturing Gentle
Monsters. Elytra and Antenna. ISBN 978-0980240122.
13. 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 (https://doi.org/10.1206%2F674.1). hdl:2246/5981 (https://hdl.handle.net/224
6%2F5981). S2CID 59942800 (https://api.semanticscholar.org/CorpusID:59942800).
14. "Giant whip scorpion Mastigoproctus giganteus giganteus (Lucas, 1835)" (http://entnemdept.ufl.
edu/creatures/misc/misc/giant_whip_scorpion.htm). Featured Creatures. Institute of Food and
Agricultural Sciences, University of Florida. Retrieved 2016-08-01.

External links
Video of vinegaroon mating behavior (https://www.youtube.com/watch?v=3Vsk6IwOOZw)

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