Palpigradi Temporal range: | |
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Eukoenenia spelaea | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Subphylum: | Chelicerata |
Class: | Arachnida |
Order: | Palpigradi Thorell, 1888 [1] |
Families | |
Synonyms | |
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Palpigradi is an order of very small arachnids commonly known as microwhip scorpion or palpigrades.
Palpigrades belong to the arachnid class. [3] They are the sister group to Solifugae, [4] no more than 3 millimetres (0.12 in) in length, [3] and averaging 1–1.5 mm (0.04–0.06 in). [5] They have a thin, pale, segmented integument, and a segmented abdomen that terminates in a whip-like flagellum. This is made up of 15 segment-like parts, or "articles", and may make up as much as half the animal's length. [6] Each article of the flagellum bears bristles, giving the whole flagellum the appearance of a bottle brush. [6] The carapace is divided into two plates between the third and fourth leg pair of legs. They have no eyes.
As in some other arachnids, the first pair of legs is modified to serve as sensory organs, and are held clear of the ground while walking. Often, however, palpigrades use their pedipalps for locomotion, so that the animal appears to be walking on five pairs of legs. [6] But they do not swing in phase with the walking legs, and are mostly used as legs in rough terrain. [7] Both the nine-segmented pedipalps and the four pairs of legs end in three claws each. The first pair of legs are 11-segmented, the second and third pairs seven-segmented and the fourth pair eight-segmented. [8] [9]
The family Prokoeneniidae have three pairs of lung-sacs on the fourth, fifth and sixth abdominal segments, although these are not true book lungs as there is no trace of the characteristic leaflike lamellae which defines book lungs. Family Eukoeneniidae have no respiratory organs at all and breathe directly through the cuticle. [10]
Their exoskeleton is very weakly sclerotized compared to other arachnids, which is the reason why fossils are so rare, and go no further back than 99 million years ago in Burmese Amber. [11]
Species of Palpigradi live interstitially in wet tropical and subtropical soils. [5] A few species have been found in shallow coral sands and on tropical beaches. [12] In Europe, they have been found in caves and underground spaces. [13] There is one endemic species on the island of Malta, in the Mediterranean Sea, which exists only in one specific cave. [3] They need a damp environment to survive, and they always hide from light, so they are commonly found in the moist earth under buried stones and rocks. They can be found on every continent, except in Arctic and Antarctic regions. Terrestrial Palpigradi have hydrophobic cuticles, but littoral (beach-dwelling) species are able to pass through the water surface easily. [12]
Very little is known about palpigrade behavior. [6] They are generally believed to be predators like their larger relatives, feeding on minuscule animals in their habitat. [6] However, their chelicerae have been described as "more like a comb or brush than the forceps of a predator", and the species Eukoenenia spelaea has been shown to feed on cyanobacteria ("blue-green algae"). [13] Their mating habits are unknown, except that they lay only a few relatively large eggs at a time. [6]
Palpigradi is split into two families, differentiated by the presence of ventral sacs on sternites IV–VI in Prokoeneniidae, and their absence in Eukoeneniidae. [14]
Two fossil palpigrade species have been described. The first one is from the Onyx Marble of Arizona, which is probably of Pliocene age. [15] Its familial position is uncertain. The second one ( Electrokoenenia yaksha ), belonging to the family Eukoeneniidae, is known from Cretaceous (Cenomanian) Burmese amber from northern Myanmar. [16] Older publications refer to a fossil palpigrade (or palpigrade-like animal) from the Jurassic of the Solnhofen limestone in Germany, [17] but this has now been shown to be a misidentified fossil insect. [18]
As of September 2022 [update] , the World Palpigradi Catalog accepts the following eight genera: [19]
The subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda. Chelicerates include the sea spiders, horseshoe crabs, and arachnids, as well as a number of extinct lineages, such as the eurypterids and chasmataspidids.
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.
Pseudoscorpions, also known as false scorpions or book scorpions, are small, scorpion-like arachnids belonging to the order Pseudoscorpiones, also known as Pseudoscorpionida or Chelonethida.
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 Thelyphonida. 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 thornlike puncture injuries.
Schizomida, also known as sprickets or short-tailed whip-scorpions, is an order of arachnids, generally less than 5 millimetres (0.20 in) in length. The order is not yet widely studied. E. O. Wilson has identified schizomids as among the "groups of organisms that desperately need experts to work on them."
The Opiliones are an order of arachnids, colloquially known as harvestmen, harvesters, harvest spiders, or daddy longlegs. As of July 2024, over 6,650 species of harvestmen have been discovered worldwide, although the total number of extant species may exceed 10,000. The order Opiliones includes five suborders: Cyphophthalmi, Eupnoi, Dyspnoi, Laniatores, and Tetrophthalmi, which were named in 2014.
Uropygi is an arachnid order comprising invertebrates commonly known as whip scorpions or vinegaroons. They are often called uropygids. 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. The order may also be called Thelyphonida. Both names, Uropygi and Thelyphonida, may be used either in a narrow sense for the order of whip scorpions, or in a broad sense which includes the order Schizomida.
Ricinulei is a small order of arachnids. Like most arachnids, they are predatory; eating small arthropods. They occur today in west-central Africa (Ricinoides) and the Americas from South America to as far north as Texas, where they either inhabit leaf-litter or caves. As of 2022, 103 extant species of ricinuleids have been described worldwide, all in the single family Ricinoididae. In older works they are sometimes referred to as Podogona. Due to their obscurity they do not have a proper common-name, though in academic literature they are occasionally referred to as hooded tickspiders.
A book lung is a type of respiration organ used for atmospheric gas-exchange that is present in many arachnids, such as scorpions and spiders. Each of these organs is located inside an open, ventral-abdominal, air-filled cavity (atrium) and connects with its surroundings through a small opening for the purpose of respiration.
The Mesothelae are a suborder of spiders. As of April 2024, two extant families were accepted by the World Spider Catalog, Liphistiidae and Heptathelidae. Alternatively, the Heptathelidae can be treated as a subfamily of a more broadly circumscribed Liphistiidae. There are also a number of extinct families.
Solifugae is an order of arachnids known variously as solifuges, sun spiders, camel spiders, and wind scorpions. The order includes more than 1,000 described species in about 147 genera. Despite the common names, they are neither true scorpions nor true spiders. Because of this, it is less ambiguous to call them "solifuges". Most species of solifuge live in dry climates and feed opportunistically on ground-dwelling arthropods and other small animals. The largest species grow to a length of 12–15 cm (5–6 in), including legs. A number of urban legends exaggerate the size and speed of solifuges, and their potential danger to humans, which is negligible.
Pulmonoscorpius is an extinct genus of scorpion from the Mississippian of Scotland. It contains a single named species, Pulmonoscorpius kirktonensis. It was one of the largest scorpions to have ever lived, with the largest known individual having an estimated length exceeding 70 cm. Pulmonoscorpius retains several general arthropod features which are absent in modern scorpions, such as large lateral eyes and a lack of adaptations for a burrowing lifestyle. It was likely an active diurnal predator, and the presence of book lungs indicate that it was fully terrestrial.
Spiders have existed since at least 380 million years. The group's origins lie within an arachnid sub-group defined by the presence of book lungs ; the arachnids as a whole evolved from aquatic chelicerate ancestors. More than 45,000 extant species have been described, organised taxonomically in 3,958 genera and 114 families. There may be more than 120,000 species. Fossil diversity rates make up a larger proportion than extant diversity would suggest with 1,593 arachnid species described out of 1,952 recognized chelicerates. Both extant and fossil species are described annually by researchers in the field. Major developments in spider evolution include the development of spinnerets and silk secretion.
Electrokoenenia yaksha is a palpigrade that lived approximately 100 million years ago during the Cretaceous period. It is the first microwhip scorpion fossil from this period to be found and is currently the oldest known Palpigrade.
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 nocturnal. At least some species of Phrynidae hold territories that they defend from other individuals.
Uraraneida is an extinct order of Paleozoic arachnids related to modern spiders. Two genera of fossils have been definitively placed in this order: Attercopus from the Devonian of United States and Permarachne from the Permian of Russia. Like spiders, they are known to have produced silk, but lack the characteristic spinnerets of modern spiders, and retain elongate telsons.
Gerontoformica is an extinct genus of stem-group ants. The genus contains thirteen described species known from Late Cretaceous fossils found in Asia and Europe. The species were described between 2004 and 2016, with a number of the species formerly being placed into the junior synonym genus Sphecomyrmodes.
Chimerarachne is a genus of extinct arachnids, containing five species. Fossils of Chimerarachne were discovered in Burmese amber from Myanmar which dates to the mid-Cretaceous, about 100 million years ago. It is thought to be closely related to spiders, but outside any living spider clade. The earliest spider fossils are from the Carboniferous, requiring at least a 170 myr ghost lineage with no fossil record. The size of the animal is quite small, being only 2.5 millimetres (0.098 in) in body length, with the tail being about 3 millimetres (0.12 in) in length. These fossils resemble spiders in having two of their key defining features: spinnerets for spinning silk, and a modified male organ on the pedipalp for transferring sperm. At the same time they retain a whip-like tail, rather like that of a whip scorpion and uraraneids. Chimerarachne is not ancestral to spiders, being much younger than the oldest spiders which are known from the Carboniferous, but it appears to be a late survivor of an extinct group which was probably very close to the origins of spiders. It suggests that there used to be spider-like animals with tails which lived alongside true spiders for at least 200 million years.
Burmese amber is fossil resin dating to the early Late Cretaceous Cenomanian age recovered from deposits in the Hukawng Valley of northern Myanmar. It is known for being one of the most diverse Cretaceous age amber paleobiotas, containing rich arthropod fossils, along with uncommon vertebrate fossils and even rare marine inclusions. A mostly complete list of all taxa described up to the end of 2023 can be found in Ross (2024).
Eukoenenia strinatii is a species of palpigrades, also known as microwhip scorpions, in the Eukoeneniidae family. The species is believed to only reside in the Cave of Bossea in Italy. It was last documented and verified in 2019.