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Palpigrades, commonly known as microwhip scorpions, are arachnids belonging to the order Palpigradi.


Description

Palpigrades belong to the
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 vinegaroon ...
class. They are the sister group to
Solifugae Solifugae is an order of animals in the class Arachnida known variously as camel spiders, wind scorpions, sun spiders, or solifuges. The order includes more than 1,000 described species in about 147 genera. Despite the common names, they are ne ...
, no more than in length, and averaging . They have a thin, pale, segmented
integument In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, shell, germ or rind. Etymology The term is derived from ''integumentum'', which is Latin for "a covering". In a transferred, or ...
, 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. Each article of the flagellum bears
bristle A bristle is a stiff hair or feather (natural or artificial), either on an animal, such as a pig, a plant, or on a tool such as a brush or broom. Synthetic types Synthetic materials such as nylon are also used to make bristles in items such as br ...
s, giving the whole flagellum the appearance of a bottle brush. 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
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") and ...
s for locomotion, so that the animal appears to be walking on five pairs of legs. But they do not swing in phase with the walking legs, and is mostly used as legs in rough terrain. Both the nine-segmented pedipalps and the four pairs of legs ends 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. The family Prokoeneniidae have three pairs of lung-sacs on the fourth, fifth and sixth abdominal segments, although these are not true
book lung 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 conn ...
s 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. Their
Exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
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.


Ecology and behavior

Species of Palpigradi live interstitially in wet
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 The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Geographical z ...
soils. A few species have been found in shallow coral sands and on tropical beaches. In Europe, they have been found in caves and underground spaces. There is one
endemic species Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
on the island of Malta, in the
Mediterranean Sea The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the ea ...
, which exists only in one specific cave. 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 In chemistry, hydrophobicity is the physical property of a molecule that is seemingly repelled from a mass of water (known as a hydrophobe). In contrast, hydrophiles are attracted to water. Hydrophobic molecules tend to be nonpolar and, th ...
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
s, but littoral (beach-dwelling) species are able to pass through the water surface easily. Very little is known about palpigrade behavior. They are generally believed to be predators like their larger relatives, feeding on minuscule animals in their habitat. However, their chelicerae have been described as "more like a comb or brush than the forceps of a predator", and the species ''
Eukoenenia spelaea ''Eukoenenia'' is a genus of Eukoeneniid microwhip scorpions, first described by Carl Julius Bernhard Börner Carl Julius Bernhard Börner (28 May 1880 – 14 June 1953) was a German entomologist. Börner was born in Bremen and died in N ...
'' has been shown to feed on
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
("blue-green algae"). Their mating habits are unknown, except that they lay only a few relatively large eggs at a time.


Classification

Palpigradi is split into two families, differentiated by the presence of ventral sacs on
sternite The sternum (pl. "sterna") is the ventral portion of a segment of an arthropod thorax or abdomen. In insects, the sterna are usually single, large sclerites, and external. However, they can sometimes be divided in two or more, in which case the ...
s IV–VI in
Prokoeneniidae Prokoeneniidae is a family of microscorpions in the order Palpigradi. There are at least two genera and about seven described species in Prokoeneniidae. Genera * '' Prokoenenia'' Borner, 1901 * ''Triadokoenenia ''Triadokoenenia'' is a monoty ...
, and their absence in
Eukoeneniidae Palpigrades, commonly known as microwhip scorpions, are arachnids belonging to the order Palpigradi. Description Palpigrades belong to the arachnid class. They are the sister group to Solifugae, no more than in length, and averaging . They have ...
. Two fossil palpigrade species have been described. The first one is from the
Onyx Marble The Onyx Marble is a geologic formation in Arizona. It preserves fossils dating back to the Neogene period. See also * List of fossiliferous stratigraphic units in Arizona * Paleontology in Arizona Paleontology in Arizona refers to paleontol ...
of
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...
, which is probably of
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Electrokoenenia yaksha ''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. The pa ...
''), belonging to the family Eukoeneniidae, is known from
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
(
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
)
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 ...
from northern
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...
. Older publications refer to a fossil palpigrade (or palpigrade-like animal) from the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
of the Solnhofen limestone in Germany, but this has now been shown to be a misidentified fossil insect.


Genera

, the World Palpigradi Catalog accepts the following eight genera: *'' Allokoenenia'' Silvestri, 1913 *'' Eukoenenia'' Börner, 1901 *''
Koeneniodes ''Koeneniodes'' is a genus of Eukoeneniidae, Eukoeneniid microwhip scorpions, first described by Filippo Silvestri in 1913. Species , the World Palpigradi Catalog accepts the following eight species: * ''Koeneniodes berndi'' Condé, 1988 ...
'' Silvestri, 1913 *'' Leptokoenenia'' Condé, 1965 *'' Prokoenenia'' Börner, 1901 *''
Triadokoenenia ''Triadokoenenia'' is a monotypic genus of Prokoeneniid microwhip scorpions, first described by Bruno Condé in 1991. Its single species, ''Triadokoenenia'' ''millotorum'' is distributed in Madagascar Madagascar (; mg, Madagasikara, ) ...
'' Condé, 1991 *†'' Electrokoenenia'' Engel & Huang, 2016 *†'' Paleokoenenia'' Rowland & Sissom, 1980


See also

* Maltese palpigrade


References

{{Taxonbar, from=Q19117 Cave arachnids Arachnid orders Cenomanian first appearances Extant Cenomanian first appearances