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Amblypygi is an
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
of
arachnids 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. Adult 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 A flagellum (; : flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores ( zoospores), and from a wide range of microorganisms to provide motility. Many pr ...
that is otherwise seen in whip-scorpions. Amblypygids possess no
silk Silk is a natural fiber, natural protein fiber, some forms of which can be weaving, woven into textiles. The protein fiber of silk is composed mainly of fibroin and is most commonly produced by certain insect larvae to form cocoon (silk), c ...
glands or
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a sti ...
. They rarely bite if threatened but can grab fingers with their
pedipalps Pedipalps (commonly shortened to palps or palpi) are the secondary pair of forward appendages among Chelicerata, chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to ...
, resulting in thorn-like puncture-injuries. As of 2023, five families, 17 genera, and around 260 species had been discovered and described. They are found in
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
and
subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones immediately to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Ge ...
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 Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
.
Fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
ized amblypygids have been found dating back to the
Carboniferous period The Carboniferous ( ) is a geologic period and system of the Paleozoic era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the Permian Period, Ma. It is the fifth and penultimate perio ...
, such as ''
Weygoldtina ''Weygoldtina'' is an extinct genus of tailless whip scorpion known from Carboniferous period, and the only known member of the family Weygoldtinidae. It is known from two species described from North America and England and originally describe ...
''.


Description


Body-plan

Being arachnids, Amblypygi possess two body-segments; the
prosoma The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...
and the
opisthosoma The opisthosoma is the posterior part of the body in some arthropods, behind the prosoma ( cephalothorax). It is a distinctive feature of the subphylum Chelicerata (arachnids, horseshoe crabs and others). Although it is similar in most respects ...
(often referred to as the
cephalothorax The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...
and abdomen), four pairs of legs, pedipalps, and
chelicerae The chelicerae () are the arthropod mouthparts, mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as "jaws", chelicerae may be shaped as either articulated ...
. Their bodies are broad and highly flattened, with a solid prosoma and a segmented opisthosoma. Amblypygids range from in legspan. Most species have eight eyes; a pair of median eyes at the front of the
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...
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.


Pedipalps

Amblypygids have
raptorial In biology (specifically the anatomy of arthropods), the term ''raptorial'' implies much the same as ''predatory'' but most often refers to modifications of an arthropod leg, arthropod's foreleg that make it function for the grasping of prey whi ...
pedipalp Pedipalps (commonly shortened to palps or palpi) are the secondary pair of forward appendages among Chelicerata, chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to ...
s modified for grabbing and retaining prey, much like the forelegs of mantisses. 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. Recent work suggests that the pedipalps display sexual dimorphism in their size and shape. 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 Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position provi ...
end of the appendage.


Exoskeleton

Whip-spiders are covered with a layer of a solidified secretion that forms a super-hydrophobic coating. 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 Tetrapulmonata is a Taxonomic rank, non-ranked Order (biology), supra-ordinal clade of arachnids. It is composed of the Extant taxon, extant orders Uropygi (whip scorpions), Schizomida (short-tailed whip scorpions), Amblypygi (tail-less whip scor ...
.


Behavior

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
pedipalp Pedipalps (commonly shortened to palps or palpi) are the secondary pair of forward appendages among Chelicerata, chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to ...
s, impaling and immobilizing the prey. This is typically done while climbing the side of a vertical surface and looking downward at their prey. Pincer-like
chelicerae The chelicerae () are the arthropod mouthparts, mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as "jaws", chelicerae may be shaped as either articulated ...
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 In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is a process by which an animal casts off parts of its body to serve some beneficial purpose, either at ...
. 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. Courtship involves the male depositing stalked
spermatophore A spermatophore, from Ancient Greek σπέρμα (''spérma''), meaning "seed", and -φόρος (''-phóros''), meaning "bearing", or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especiall ...
s, which have one or more sperm masses at the tip, onto the ground, and using his pedipalps to guide the female over them. She gathers the sperm and lays fertilized
eggs An egg is an organic vessel in which an embryo begins to develop. Egg, EGG or eggs may also refer to: Biology * Egg cell, the female reproductive cell (gamete) in oogamous organisms Food * Eggs as food Places * Egg, Austria * Egg, Switzerland ...
into a sac carried under the
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
, or
opisthosoma The opisthosoma is the posterior part of the body in some arthropods, behind the prosoma ( cephalothorax). It is a distinctive feature of the subphylum Chelicerata (arachnids, horseshoe crabs and others). Although it is similar in most respects ...
. 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 Cornell University is a Private university, private Ivy League research university based in Ithaca, New York, United States. The university was co-founded by American philanthropist Ezra Cornell and historian and educator Andrew Dickson W ...
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. Amblypygids hold territories that they defend from other individuals. The amblypygid diet mostly consists of arthropod prey, but these opportunistic predators have also been observed feeding on vertebrates. 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. Molting is done while hanging from the underside of a horizontal surface in order to use
gravity In physics, gravity (), also known as gravitation or a gravitational interaction, is a fundamental interaction, a mutual attraction between all massive particles. On Earth, gravity takes a slightly different meaning: the observed force b ...
to assist in separating the old exoskeleton from the animal.


As pets

Several genera of Amblypygi are sold and kept as
pet A pet, or companion animal, is an animal kept primarily for a person's company or entertainment rather than as a working animal, livestock, or a laboratory animal. Popular pets are often considered to have attractive/ cute appearances, inte ...
s, including '' Acanthophrynus'', '' Charinus,'' ''
Charon In Greek mythology, Charon or Kharon ( ; ) is a psychopomp, the ferryman of the Greek underworld. He carries the souls of those who have been given funeral rites across the rivers Acheron and Styx, which separate the worlds of the living and ...
,'' '' Damon, Euphrynichus,'' '' Heterophrynus'', ''
Phrynus ''Phrynus'' is a genus of whip spiders found in tropical and subtropical regions, mostly in the new world. Appearance Like other species of the order Amblypygi, species of the genus ''Phrynus'' are dorso-ventrally flattened arachnids with elonga ...
'', ''Paraphrynus'', and ''Phrynichus''. 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 and . 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.


Genera

The following genera are recognised: * Palaeoamblypygi Weygoldt, 1996 **
Paracharontidae Paracharontidae is an arachnid family within the order Amblypygi (tailless whip scorpions). Paracharontidae and the extinct Weygoldtinidae from the Carboniferous form the suborder Paleoamblypygi, the sister group to the remaining Amblypygi. The ...
Weygoldt, 1996 *** ''
Paracharon ''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 Ame ...
'' Hansen, 1921 (1 species, West Africa) *** '' Jorottui'' Moreno-González, Gutierrez-Estrada, & Prendini, 2023 (1 species, northern South America) **
Weygoldtinidae ''Weygoldtina'' is an extinct genus of Amblypygi, tailless whip scorpion known from Carboniferous period, and the only known member of the family Weygoldtinidae. It is known from two species described from North America and England and originall ...
Dunlop, 2018 *** †''
Weygoldtina ''Weygoldtina'' is an extinct genus of tailless whip scorpion known from Carboniferous period, and the only known member of the family Weygoldtinidae. It is known from two species described from North America and England and originally describe ...
'' Dunlop, 2018 (2 species, Upper Carboniferous Europe, North America) * Euamblypygi Weygoldt, 1996 ** †'' Paracharonopsis'' Engel & Grimaldi, 2014 (1 species,
Cambay amber Khambhat state or Cambay state was a princely state in India during the British Raj. The city of Khambhat in present-day Gujarat was its capital. The state was bounded in the north by the Kaira district (Kheda) and in the south by the Gulf ...
, India,
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
) **
Charinidae Charinidae is an arachnid Arachnids are arthropods in the Class (biology), class Arachnida () of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, Solifugae, ...
Weygoldt, 1996 *** '' Charinus'' Simon, 1892 (33 species) ***'' Sarax'' Simon, 1892 (10 species) ***'' Weygoldtia'' Miranda, Giupponi, Prendini & Scharff, 2018 (3 species) **Neoamblypygi Weygoldt, 1996 ***
Charontidae ''Charon'' is a genus of whip-spiders from Asia and Australasia, first described by Ferdinand Karsch Ferdinand Anton Franz Karsch (2 September 1853, in Münster – 20 December 1936, in Berlin) was a German arachnologist, entomologist and a ...
Simon, 1892 ****'' Catageus'' Thorell, 1889 (9 species) ****''
Charon In Greek mythology, Charon or Kharon ( ; ) is a psychopomp, the ferryman of the Greek underworld. He carries the souls of those who have been given funeral rites across the rivers Acheron and Styx, which separate the worlds of the living and ...
'' Karsch, 1879 (5 species) *** Unidistitarsata Engel & Grimaldi, 2014 ****†'' Kronocharon'' Engel & Grimaldi, 2014 (1 species,
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. Th ...
, Myanmar, Cretaceous) ****†'' Burmacharon''? Hu et al. 2020 (1 species, Burmese amber, Myanmar, Cretaceous) ****Phrynoidea Blanchard, 1852 *****
Phrynichidae Phrynichidae is a family of arachnid Arachnids are arthropods in the Class (biology), class Arachnida () of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, ...
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 Phrynidae is a Family (biology), family of amblypygid chelicerata, arachnida arthropods also known as whip spiders and tailless whip scorpions. Phrynidae species are found in tropics, tropical and subtropics, subtropical regions in North and Sout ...
Blanchard, 1852 ******'' Acanthophrynus'' Kraepelin, 1899 (1 species) ******†'' Britopygus'' Dunlop & Martill, 2002 (1 species;
Crato Formation The Crato Formation is a geologic formation (stratigraphy), formation of Early Cretaceous (Aptian) age in northeastern Brazil's Araripe Basin. It is an important Lagerstätten, Lagerstätte (undisturbed fossil accumulation) for palaeontology, pa ...
, Brazil, Cretaceous) ******'' Heterophrynus'' Pocock, 1894 (14 species) ******''
Paraphrynus ''Paraphrynus'' is a genus of whip spiders, also known as tailless whip scorpions (order Amblypygi), of the family Phrynidae. It is distributed from the southwestern United States to Central America, including several Caribbean islands. Most spec ...
'' Moreno, 1940 (18 species) ******''
Phrynus ''Phrynus'' is a genus of whip spiders found in tropical and subtropical regions, mostly in the new world. Appearance Like other species of the order Amblypygi, species of the genus ''Phrynus'' are dorso-ventrally flattened arachnids with elonga ...
'' Lamarck, 1801 (28 species, Oligocene - Recent) *''
Incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
:'' ** † '' Sorellophrynus'' Harvey, 2002 (1 species, Upper Carboniferous, North America) ** † '' Thelyphrynus'' Petrunkevich, 1913 (1 species, Upper Carboniferous, North America)


References


External links

*
Amblypigid video
summarizing research from University of Nebraska's Eben Gering * {{Authority control Arachnid orders Amblypygi Extant Pennsylvanian first appearances Carboniferous arachnids Permian arachnids Mesozoic arachnids Cenozoic arachnids