Amblypygi
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Amblypygi is an ancient order of
arachnid 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 vinegar ...
chelicerate
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s also known as African cave-dwelling spiders, whip spiders and tailless whip scorpions (not to be confused with
whip scorpion 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 ...
s 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 (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
that is otherwise seen in whip scorpions. Amblypygids possess no
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the ...
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 st ...
ous fangs. They rarely bite if threatened, but can grab fingers with their
pedipalps Pedipalps (commonly shortened to palps or palpi) 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 ...
, resulting in thorn-like puncture injuries. As of 2016, 5 families, 17 genera and around 155 species had been discovered and described. They are found in
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
and subtropical regions worldwide; they are mainly found in warm and humid environments and like to stay protected and hidden within leaf litter, caves, or underneath bark. Some species are subterranean; all are nocturnal.
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 preserved ...
ized amblypygids have been found dating back to the
Carboniferous period The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
, such as ''
Weygoldtina ''Weygoldtina'' is an extinct genus of tailless whip scorpion known from Carboniferous fossil record, and the only known member of the family Weygoldtinidae. The genus is known from two species described from North America and England and origi ...
''.


Description

Amblypygids range from in legspan. Their bodies are broad and highly flattened, with a solid carapace and a segmented
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
, 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 to ...
. 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. Amblypygids have raptorial
pedipalp Pedipalps (commonly shortened to palps or palpi) 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") an ...
s modified for grabbing and retaining prey, much like those of a
mantis Mantises are an order (Mantodea) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate and tropical habitats. They ha ...
; recent work suggests that the pedipalps display sexual dimorphism in their size and shape. The first pair of legs act as sensory organs and are not used for walking. The sensory legs are very thin and elongate, have numerous sensory receptors, and can extend several times the length of the body.


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 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") an ...
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 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 fangs, or similarl ...
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 premolt. 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 or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophores ...
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 Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
into a sac carried under the
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
, 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 to ...
. 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 statutory land-grant research university based in Ithaca, New York. It is a member of the Ivy League. Founded in 1865 by Ezra Cornell and Andrew Dickson White, Cornell was founded with the intention to tea ...
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 before, during, and after molting. Like other arachnids, an amblypygid will molt several times during its life. Molting is done from hanging from the underside of a horizontal surface in order to use
gravity In physics, gravity () is a fundamental interaction which causes mutual attraction between all things with mass or energy. Gravity is, by far, the weakest of the four fundamental interactions, approximately 1038 times weaker than the stro ...
to assist in separating the old exoskeleton from the animal.


As pets

Several species 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 appearances, intelligence ...
s including '' Damon diadema'', ''
Damon medius ''Damon medius'' is a species of arachnid of the family Phrynichidae. Description ''Damon medius'' can reach a body length of . This strictly arboreal and nocturnal whip spider has a flat body and fragile legs, by which it moves sideways like a ...
'', '' Damon variegatus'', '' Euphrynichus amanica'', '' Heterophrynus batesii'', '' Acanthophrynus coronatus'', '' Phrynus marginemaculatus'' and '' Paraphrynus mexicanus''. Tailless whip scorpions are kept in tall (>45 cm; >18 in) glass enclosures which allow for two things: Enough vertical space for climbing and moulting, and enough space for heat to dissipate in order to keep the enclosure between 70 °F and 75 °F. Five centimetres (two inches) 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 5–10 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 of tailless whip scorpions. Paracharontidae and Weygoldtina, Weygoldtinidae form the suborder Paleoamblypygi, the putative sister group to the remaining Amblypygi; this suborder is primarily ...
Weygoldt, 1996 *'' Paracharon'' Hansen, 1921 (1 described species +1 undescribed species, West Africa, northern South America) *†'' Paracharonopsis'' Engel & Grimaldi, 2014 (1 species,
Cambay amber Cambay, Kambay or Khambhat was a princely state in India during the British Raj. The City of Khambat (Cambay) in present-day Gujarat was its capital. The state was bounded in the north by the Kaira district and in the south by the Gulf of C ...
, India,
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
) ; Weygoldtinidae Dunlop, 2018 *†''
Weygoldtina ''Weygoldtina'' is an extinct genus of tailless whip scorpion known from Carboniferous fossil record, and the only known member of the family Weygoldtinidae. The genus is known from two species described from North America and England and origi ...
'' Dunlop, 2018 (2 species, Carboniferous) ; Euamblypygi Weygoldt, 1996 ;
Charinidae Charinidae is an arachnid family within the order of tailless whip scorpions.Harvey, M.S. 2003. Order Amblypygi. pp. 59–99 in, Catalogue of the Smaller Arachnid Orders of the World: Amblypygi, Uropygi, Schizomida, Palpigradi, Ricinulei and Soli ...
Weygoldt, 1996 *''
Charinus ''Charinus'' is a genus of amblypygids of the family Charinidae Charinidae is an arachnid family within the order of tailless whip scorpions.Harvey, M.S. 2003. Order Amblypygi. pp. 59–99 in, Catalogue of the Smaller Arachnid Orders of the Wo ...
'' Simon, 1892 (33 species) *'' Sarax'' Simon, 1892 (10 species) *'' Weygoldtia'' Miranda, Giupponi, Prendini & Scharff, 2018 (3 species) ; Neoamblypygi Weygoldt, 1996 ;
Charontidae Charontidae is a family of whip spiders. Genera *''Catageus'' Thorell, 1889 (9 species) *''Charon In Greek mythology, Charon or Kharon (; grc, Χάρων) is a psychopomp, the ferryman of Hades, the Greek underworld. He carries the so ...
Simon, 1892 *''
Catageus ''Catageus'' is a genus of amblypygids of the family Charontidae. Species , there are 8 species in this genus. * ''Catageus berkeleyi'' (Frederic Henry Gravely, Graveley, 1915) * ''Catageus brevispina'' (Weygoldt, 2002) * ''Catageus cavernicola ...
'' Thorell, 1889 (9 species) *'' Charon'' Karsch, 1879 (5 species) ; Unidistitarsata Engel & Grimaldi, 2014 family unspecified ;*†'' 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. The ...
, Myanmar, Cretaceous) ; Phrynoidea Blanchard, 1852 ;
Phrynichidae Phrynichidae is a family of arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, c ...
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 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 no ...
Blanchard, 1852 *''
Acanthophrynus ''Acanthophrynus'' is a genus of tailless whipscorpions in the family Phrynidae. There is at least one described species in ''Acanthophrynus'', ''A. coronatus''. ''Acanthophrynus coronatus'' ''Acanthophrynus coronatus'' is a tailless whipsco ...
'' Kraepelin, 1899 (1 species) *†'' Britopygus'' Dunlop & Martill, 2002 (1 species;
Crato Formation The Crato Formation is a geologic formation of Early Cretaceous (Aptian) age in northeastern Brazil's Araripe Basin. It is an important Lagerstätte (undisturbed fossil accumulation) for palaeontologists. The strata were laid down mostly during ...
, Brazil, Cretaceous) *†'' Electrophrynus'' Petrunkevich, 1971 (1 species; Mexican amber, Miocene) *''
Heterophrynus ''Heterophrynus'' is a genus of whip spiders, also known as tailless whip scorpions ( order Amblypygi), of the family Phrynidae. It is mostly distributed in South America. Species There are 16 species: *'' Heterophrynus alces'' Pocock, 1902 *'' ...
'' 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 sp ...
'' 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 elon ...
'' Lamarck, 1801 (28 species, Oligocene - Recent) ;''
incertae sedis ' () or ''problematica'' is a term used for a 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, uncertain ...
'' * † '' Sorellophrynus'' Harvey, 2002 (1 species, Upper Carboniferous) * † '' Thelyphrynus'' Petrunkevich, 1913 (1 species, Upper Carboniferous)


References


External links

*
Amblypigid video
summarizing research from University of Nebraska's Eben Gering

{{Taxonbar, from=Q19111 Arachnid orders Extant Pennsylvanian first appearances Prehistoric arthropod orders Carboniferous arachnids Permian arachnids Mesozoic arachnids Cenozoic arachnids