Amblypygi

{{Short description|Order of arachnids}} {{Distinguish|Ariamnes colubrinus}} {{Automatic taxobox | fossil_range = Bashkirian/Moscovian–Holocene {{fossil_range|315|0|ref=<ref name="Garwood2017">{{cite journal | last1 = Garwood | first1 = Russell J. | last2 = Dunlop | first2 = Jason A. | last3 = Knecht | first3 = Brian J. | last4 = Hegna | first4 = Thomas A. | date = 2017 | title = The phylogeny of fossil whip spiders | journal = BMC Evolutionary Biology | volume = 17 | issue = 1| page = 105 | doi = 10.1186/s12862-017-0931-1 | pmid=28431496 | pmc=5399839 | bibcode = 2017BMCEE..17..105G | doi-access = free }}</ref>}} | image = Heterophrynus 02.jpg | image_caption = ''Heterophrynus'', Ecuador | taxon = Amblypygi | authority = Thorell, 1883 | subdivision_ranks = Families | subdivision = *'''Palaeoamblypygi''' **Paracharontidae **{{extinct}}Weygoldtinidae *'''Euamblypygi''' **Charinidae **Charontidae **Phrynichidae **Phrynidae | range_map = Amblypygi Distribution Map.png | synonyms = *Phrynéides {{small|Walckenaer, 1837}} *Phrynichida {{small|Petrunkevitch, 1945}} | synonyms_ref = <ref>{{cite web | url = https://museum.wa.gov.au/catalogues-beta/whip-spiders/order/amblypygi | title = Amblypygi {{small|Thorell 1883}} | first = M.S. | last = Harvey | date = 2013 | work = Whip spiders of the World | publisher = Western Australian Museum | version = version 1.0. | access-date = 23 September 2024 | archive-url = https://web.archive.org/web/20240923192643/https://museum.wa.gov.au/catalogues-beta/whip-spiders/order/amblypygi | archive-date = 23 September 2024}}</ref> }}

'''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 Uropygi. 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.

{{As of|2026}}, five families, 18 genera, and around 280 species had been discovered and described.<ref>{{cite web |date=2023 |title=World Amblypygi Catalog |url=https://wac.nmbe.ch/order/amblypygi/1 |access-date=17 April 2026 |website=World Amblypygi Catalog |publisher=Natural History Museum Bern}}</ref> They are found in tropical and subtropical regions worldwide, mainly in warm and humid environments. They like to stay protected and hidden within leaf litter, caves, or underneath bark. Some species are subterranean; all are nocturnal. Fossilized amblypygids have been found dating back to the Carboniferous period, such as ''Weygoldtina''.<ref name="Garwood2017" />

==Description== thumb|Detail of pedipalps thumb|right|Parts of an amblypygid, from Pocock (1900)<ref>{{cite book|author=R. I. Pocok|year=1900|title=Fauna of British India. Arachnida|url=http://www.ub.ntnu.no/scorpion-files/pocock_fauna_india.php|author-link=Reginald Innes Pocock}}</ref>

===Body-plan=== Being arachnids, Amblypygi possess two body-segments; the prosoma and the opisthosoma (often referred to as the cephalothorax and abdomen), four pairs of legs, pedipalps, and chelicerae. Their bodies are broad and highly flattened, with a solid prosoma and a segmented opisthosoma.<ref name=":1" />

Amblypygids range from {{convert|5|to|16|cm|sp=us}} in walking legspan.<ref name="Chapin">{{cite journal |last1=Chapin |first1=KJ |last2=Hebets |first2=EA |year=2016 |title=Behavioral ecology of amblypygids |url=http://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1056&context=bioscihebets |journal=Journal of Arachnology |volume=44 |issue=1 |pages=1–14 |doi=10.1636/V15-62.1 |s2cid=29923727|url-access=subscription }}</ref><ref>{{Cite book |last1=Weygoldt |first1=Peter |title=Whip Spiders (Chelicerata: Amblypygi): Their Biology, Morphology and Systematics |publisher=Apollo Books |year=2000 |isbn=8788757463}}</ref> Most species have eight eyes; a pair of median eyes at the front of the carapace above the chelicerae and 2 smaller clusters of three eyes each further back on each side.

The first pair of legs act as sensory organs and are not used for walking. The sensory legs are very thin and elongated, have numerous sensory receptors, and can extend several times the length of the body.<ref name="Chapin" /><ref name=":1" />

====Pedipalps==== Amblypygids have raptorial pedipalps modified for grabbing and retaining prey, much like the forelegs of mantisses.<ref name="IZ">{{cite book |author=Robert D. Barnes |year=1982 |title= Invertebrate Zoology |publisher=Holt-Saunders International |location= Philadelphia, PA |pages= 617–619|isbn= 0-03-056747-5}}</ref> The pedipalps are generally covered in spines, used for impaling and capturing prey. They are kept folded in front of the prosoma when not in use.<ref name=":1" /> Recent work suggests that the pedipalps display sexual dimorphism in their size and shape.<ref name="McLeanGarwood2019">{{cite journal|last1=McLean|first1=C.J.|last2=Garwood|first2=R.J.|last3=Brassey|first3=C.A.|title=Sexual dimorphism in the size and shape of the raptorial pedipalps of Giant Whip Spiders (Arachnida: Amblypygi)|journal=Journal of Zoology|volume=310|issue=1|year=2019|pages=45–54|issn=0952-8369|doi=10.1111/jzo.12726|doi-access=free}}</ref>

Pedipalp anatomy varies strongly with species, with configurations often conforming to a particular style of prey capture. The pedipalps of some genera such as Euphrynicus are extremely long, and free of spines until near the extreme distal end of the appendage.<ref name=":1" />

====Exoskeleton==== Whip-spiders are covered with a layer of a solidified secretion that forms a super-hydrophobic coating.<ref>[https://zoologicalletters.biomedcentral.com/articles/10.1186/s40851-016-0059-y Whip spiders (Amblypygi) become water-repellent by a colloidal secretion that self-assembles into hierarchical microstructures]</ref> Studies on the spotted tailless whip-scorpion also show their exoskeleton is enriched with several trace-elements, including calcium, magnesium, manganese, potassium, sodium, and zinc, which tends to accumulate as the individual gets older. The same trace-elements are also present in the exoskeleton of the other members of Tetrapulmonata.<ref>[https://bioone.org/journals/the-journal-of-arachnology/volume-49/issue-2/JoA-S-20-048/Elemental-enrichment-of-the-exoskeleton-of-the-whip-spider-Phrynus/10.1636/JoA-S-20-048.short Elemental enrichment of the exoskeleton of the whip spider Phrynus marginemaculatus (Arachnida: Amblypygi)]</ref>

==Behavior== thumb|A ''Damon diadema'' mother carrying young Amblypygids have eight legs, but use only six for walking, often in a crab-like, sideways fashion. The front pair of legs are modified for use as antennae-like feelers, with many fine segments giving the appearance of a "whip". When a suitable prey is located with the antenniform legs, the amblypygid seizes its victim with large spines on the grasping pedipalps, impaling and immobilizing the prey. This is typically done while climbing the side of a vertical surface and looking downward at their prey.<ref name="Ladle2003">{{Cite journal |last1=Ladle |first1=Richard J. |last2=Velander |first2=Kathryn |date=2003 |title=Fishing behavior in a giant whip spider |url=https://www.researchgate.net/publication/289037904 |journal=The Journal of Arachnology |volume=31 |pages=154–156 |via=ResearchGate|doi=10.1636/0161-8202(2003)031[0154:FBIAGW]2.0.CO;2 |s2cid=86012520 }}</ref>

Pincer-like chelicerae then work to grind and chew the prey prior to ingestion. The tailless whip scorpion may go for over a month in which no food is eaten. Often this is due to pre-molt. Due to the lack of venom the tailless whip scorpion is very nervous in temperament, retreating away if any dangerous threat is sensed by the animal.{{Citation needed|reason=Sounds like speculation|date=March 2022}}

thumb|Comparing the front and back legs of an amblypygid Courtship involves the male depositing stalked spermatophores, which have one or more sperm masses at the tip, onto the ground, and using his pedipalps to guide the female over them.<ref>{{Cite journal|author=Peter Weygoldt |author-link=Peter Weygoldt |year=1999 |title=Spermatophores and the evolution of female genitalia in whip spiders (Chelicerata, Amblypygi) |journal=Journal of Arachnology |volume=27 |issue=1 |pages=103–116 |url=http://www.americanarachnology.org/JoA_Congress/JoA_v27_n1/arac_27_01_0103.pdf |url-status=dead |archive-url=https://web.archive.org/web/20080517002449/http://www.americanarachnology.org/JoA_Congress/JoA_v27_n1/arac_27_01_0103.pdf |archive-date=2008-05-17 }}</ref> She gathers the sperm and lays fertilized eggs into a sac carried under the abdomen, or opisthosoma. When the young hatch, they climb up onto the mother's back; any which fall off before their first molt will not survive.

Some species of amblypygids, particularly ''Phrynus marginemaculatus'' and ''Damon diadema'', may be among the few examples of arachnids that exhibit social behavior. Research conducted at Cornell University suggests that mother amblypygids communicate with their young with her antenniform front-legs, and the offspring reciprocate both with their mother and siblings. The ultimate function of this social behavior remains unknown.<ref>{{cite journal|last1=Rayor|first1=Linda|title=Social Behavior in Amblypygids, and a Reassessment of Arachnid Social Patterns|journal=Journal of Arachnology|date=December 2017|volume=31|issue=12|pages=399–421|doi=10.1636/S04-23.1|s2cid=34165769|url=https://www.biodiversitylibrary.org/part/228950 }}</ref> Amblypygids hold territories that they defend from other individuals.<ref>{{Cite journal | author1=Chapin KJ | author2=Hill-Lindsay S | year=2015 | title=Territoriality evidenced by asymmetric intruder-holder motivation in an amblypygid | journal=Behavioural Processes | volume=122 | pages=110–115| doi=10.1016/j.beproc.2015.11.014 | pmid=26616673 | s2cid=37584495 }}</ref>

The amblypygid diet mostly consists of arthropod prey, but these opportunistic predators have also been observed feeding on vertebrates.<ref name=Chapin /> Amblypygids generally do not feed for a period of time before, during, and after molting. Like other arachnids, an amblypygid will molt several times during its life.<ref name=Chapin /> Molting is done while hanging from the underside of a horizontal surface in order to use gravity to assist in separating the old exoskeleton from the animal.

==As pets== Several genera of Amblypygi are sold and kept as pets, including ''Acanthophrynus'', ''Charinus,'' ''Charon,'' ''Damon, Euphrynichus,'' ''Heterophrynus'', ''Phrynus'', ''Paraphrynus'', and ''Phrynichus''.<ref name="exotic-pets">{{cite web|url=https://www.exotic-pets.co.uk/tail-less-whip-scorpion.html |title=Tail-less Whip Scorpion - Damon medius |publisher=exotic-pets.co.uk|access-date=2019-07-13}}</ref><ref name=":1">{{Cite book |last=McMonigle |first=Orin |title=Breeding the world's largest living arachnid: amblypygid biology, natural history, and captive husbandry |date=2013 |publisher=Coachwhip Publications |isbn=978-1-61646-183-6 |location=Greenville, Ohio |pages=}}</ref> Tailless whip-scorpions are kept in tall enclosures with arboreal climbing surfaces to allow for two things: Enough vertical space for climbing and molting, and enough space for heat to dissipate in order to keep the enclosure between {{convert|70|F|C}} and {{convert|75|F|C}}. {{convert|5|cm|in|sp=us}} of substrate at the bottom of the enclosure is generally sufficient to allow for burrowing and also serves as a method to retain water in order to keep the humidity above 75%. Tailless whip-scorpions live anywhere between 5 and 10 years. Feeding can include small insects such as crickets, mealworms, and roaches.<ref>{{cite web |title=Tailless Whip Scorpion Care Sheet |url=https://www.reptilecentre.com/info-tailless-whip-scorpion-care-sheet |website=Reptile Centre |access-date=December 27, 2019}}</ref><ref>{{cite web |title=Tailless Whip Scorpion: Facts, Lifespan, Care, Feeding, & Breeding |url=https://allpetsdirectory.com/tailless-whip-scorpion/ |website=AllPetsDirectory |access-date=27 December 2019}}</ref>

==Genera== thumb|An amblypygid molting [[File:Damon johnstoni – Lydekker, 1879.png|thumb|''Damon johnstonii'' from West Africa]] The following genera are recognised:<ref name="Harvey">{{cite book |author=Mark S. Harvey |year=2003 |title=Catalogue of the smaller arachnid orders of the world: Amblypygi, Uropygi, Schizomida, Palpigradi, Ricinulei and Solifugae |publisher=CSIRO Publishing |isbn=978-0-643-06805-6 |chapter=Order Amblypygi |pages=3–58 |chapter-url=https://books.google.com/books?id=I5JNqiORopUC&pg=PA12}}</ref><ref name="Engel and Grimaldi">{{cite journal |last1=Engel|first1=M.S. |last2=Grimaldi |first2=D.A. |year=2014 |title=Whipspiders (Arachnida: Amblypygi) in amber from the Early Eocene and mid-Cretaceous, including maternal care |journal= Novitates Paleoentomologicae |volume=9 |pages=1–17}}</ref> * Palaeoamblypygi Weygoldt, 1996 ** Paracharontidae Weygoldt, 1996 *** ''Paracharon'' Hansen, 1921 (1 species, West Africa) *** ''Jorottui'' Moreno-González, Gutierrez-Estrada, & Prendini, 2023 (1 species, northern South America) ** Weygoldtinidae Dunlop, 2018<ref name=":0">{{Cite journal |last=Dunlop |first=Jason A. |date=2018-03-01 |title=Systematics of the Coal Measures whip spiders (Arachnida: Amblypygi) |url=https://www.sciencedirect.com/science/article/pii/S0044523117300967 |journal=Zoologischer Anzeiger |series=In honor of Peter Weygoldt |language=en |volume=273 |pages=14–22 |doi=10.1016/j.jcz.2017.11.004 |issn=0044-5231|url-access=subscription }}</ref> *** †''Weygoldtina'' Dunlop, 2018 (2 species, Upper Carboniferous Europe, North America)<ref name=":0" /> * Euamblypygi Weygoldt, 1996 ** †''Paracharonopsis'' Engel & Grimaldi, 2014 (1 species, Cambay amber, India, Eocene)<ref name=":02">{{Cite journal |last1=Moreno-González |first1=Jairo A. |last2=Gutierrez-Estrada |first2=Miguel |last3=Prendini |first3=Lorenzo |date=2023-06-28 |title=Systematic Revision of the Whip Spider Family Paracharontidae (Arachnida: Amblypygi) with Description of a New Troglobitic Genus and Species from Colombia |url=https://bioone.org/journals/american-museum-novitates/volume-2023/issue-4000/4000.1/Systematic-Revision-of-the-Whip-Spider-Family-Paracharontidae-Arachnida/10.1206/4000.1.full |journal=American Museum Novitates |issue=4000 |pages=1–36 |doi=10.1206/4000.1 |s2cid=259275494 |issn=0003-0082}}</ref> ** Charinidae Weygoldt, 1996 *** ''Charinus'' Simon, 1892 (33 species) ***''Sarax'' Simon, 1892 (10 species) ***''Weygoldtia'' Miranda, Giupponi, Prendini & Scharff, 2018 (3 species) **Neoamblypygi Weygoldt, 1996 ***Charontidae Simon, 1892 ****''Catageus'' Thorell, 1889 (9 species) ****''Charon'' Karsch, 1879 (5 species) ***Unidistitarsata Engel & Grimaldi, 2014 ****†''Kronocharon'' Engel & Grimaldi, 2014 (1 species, Burmese amber, Myanmar, Cretaceous) ****†''Burmacharon''? Hu et al. 2020 (1 species, Burmese amber, Myanmar, Cretaceous) ****Phrynoidea Blanchard, 1852 *****Phrynichidae Simon, 1900 ******''Damon'' C. L. Koch, 1850 (10 species) ******''Euphrynichus'' Weygoldt, 1995 (2 species) ******''Musicodamon'' Fage, 1939 (1 species) ******''Phrynichodamon'' Weygoldt, 1996 (1 species) ******''Phrynichus'' Karsch, 1879 (16 species) ******''Trichodamon'' Mello-Leitão, 1935 (2 species) ******''Xerophrynus'' Weygoldt, 1996 (1 species) *****Phrynidae Blanchard, 1852 ******''Acanthophrynus'' Kraepelin, 1899 (1 species) ******†''Britopygus'' Dunlop & Martill, 2002 (1 species; Crato Formation, Brazil, Cretaceous) ******''Heterophrynus'' Pocock, 1894 (14 species) ******''Paraphrynus'' Moreno, 1940 (18 species) ******''Phrynus'' Lamarck, 1801 (28 species, Oligocene - Recent) *''Incertae sedis:'' ** † ''Sorellophrynus'' Harvey, 2002 (1 species, Upper Carboniferous, North America) ** † ''Thelyphrynus'' Petrunkevich, 1913 (1 species, Upper Carboniferous, North America)

==References== {{Reflist|2}}

==External links== {{Commons category}} {{Portal|Arthropods}} * {{Wikispecies-inline|Amblypygi}} * [https://web.archive.org/web/20120829192507/http://www.thewildclassroom.com/biomes/speciesprofile/species/amblypigid.html Amblypigid video] summarizing research from University of Nebraska's Eben Gering * {{usurped|1=[https://web.archive.org/web/20160304030218/http://www.scorpionsetc.com/amblypygi-the-antillean-west-indian-fauna.html Amblypygi. The Antillean (West Indian) fauna.]}}

Category:Arachnid orders Category:Amblypygi Category:Extant Pennsylvanian first appearances Category:Carboniferous arachnids Category:Permian arachnids Category:Mesozoic arachnids Category:Cenozoic arachnids Category:Taxa named by Tamerlan Thorell