Mesothelae Genera
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The Mesothelae are a suborder of
spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s (order Araneae) that includes a single extant family,
Liphistiidae The spider family Liphistiidae, recognized by Tamerlan Thorell in 1869, comprises 8 genera and about 100 species of medium-sized spiders from Southeast Asia, China, and Japan. They are among the most basal living spiders, belonging to the subord ...
, and a number of extinct families. This suborder is thought to form the sister group to all other living spiders, and to retain ancestral characters, such as a segmented abdomen with spinnerets in the middle and two pairs of book lungs. Members of Liphistiidae are medium to large spiders with eight eyes grouped on a tubercle. They are found only in China, Japan, and southeast Asia. The oldest known Mesothelae spiders are known from the
Carboniferous 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 ...
, over 300 million years ago. The Heptathelidae were once considered their own family; today they are considered a subfamily of the Liphistiidae.


Description

Members of Mesothelae have paraxial
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 similarly ...
, two pairs of
coxal gland The coxal gland is a gland found in some arthropods, for collecting and excreting urine. They are found in all arachnids (with the exception of some Acari), and in other chelicerates, such as horseshoe crabs. The coxal gland is thought to be homo ...
s on the legs, eight eyes grouped on a nodule, two pairs of
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, and no endites on the base of the
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 ...
. Most have at least seven or eight
spinnerets A spinneret is a silk-spinning organ of a spider or the larva of an insect. Some adult insects also have spinnerets, such as those borne on the forelegs of Embioptera. Spinnerets are usually on the underside of a spider's opisthosoma, and are ...
near the middle of the abdomen. Lateral spinnerets are multi-segmented. Recent Mesothelae are characterized by the narrow
sternum The sternum or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major blood vessels from injury. Sh ...
on the ventral side of 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 ...
. Several
plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...
characteristics may be useful in recognizing these spiders: there are
tergite A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'mar ...
plates on the
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal co ...
side and the almost
median In statistics and probability theory, the median is the value separating the higher half from the lower half of a data sample, a population, or a probability distribution. For a data set, it may be thought of as "the middle" value. The basic fe ...
position of the spinnerets on the
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Th ...
side of 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 to a ...
. Although it has been claimed that they lack venom glands and ducts, which almost all other spiders have, subsequent works have demonstrated that at least some, possibly all, do in fact have both the glands and ducts. All Mesothelae have eight spinnerets in four pairs. Like mygalomorph spiders, they have two pairs of book lungs. Unlike all other extant mesothelians, heptathelines do not have fishing lines in front of the entrances to the burrows that they construct, making them more difficult to find. They also have a paired receptaculum (unpaired in other liphistiids), and have a conductor in their palpal organ. These long palps can confusingly look like an extra pair of legs, a mistake also made of some
solifugids 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 nei ...
.


Taxonomy

Reginald Innes Pocock Reginald Innes Pocock F.R.S. (4 March 1863 – 9 August 1947) was a British zoologist. Pocock was born in Clifton, Bristol, the fourth son of Rev. Nicholas Pocock and Edith Prichard. He began showing interest in natural history at St. Edward ...
in 1892 was the first to realize that the exceptional characters of the genus ''
Liphistius ''Liphistius'' is a genus of basal trapdoor spiders in the family Liphistiidae. They are found in Japan, China, and Southeast Asia. Etymology ''Liphistius'' is from the Greek ''leipo'' (lacking) and ''stios'' (equality). Biology Female body ...
'' (the only member of the group then known) meant that it was more different from the remaining spiders than they were among themselves. Accordingly, he proposed dividing spiders into two subgroups, Mesothelae for ''Liphistius'', and
Opisthothelae The Opisthothelae are spiders within the order Araneae, consisting of the Mygalomorphae and the Araneomorphae, but excluding the Mesothelae. The Opisthothelae are sometimes presented as an unranked clade and sometimes as a suborder of the Arane ...
for all other spiders. The names refer to the position of the spinning organs, which are in the middle of the abdomen in ''Liphistius'' and nearer the end in all other spiders. In
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
, μέσος (''mesos'') means "middle", and θήλα (''thēla'') "teat".


Phylogeny and classification

Pocock divided his Opisthothelae into two groups, which he called
Mygalomorphae The Mygalomorphae, or mygalomorphs, are an infraorder of spiders, and comprise one of three major groups of living spiders with over 3000 species, found on all continents except Antarctica. Many members are known as trapdoor spiders due to the ...
and Arachnomorphae (now
Araneomorphae The Araneomorphae (also called the Labidognatha) are an infraorder of spiders. They are distinguishable by chelicerae (fangs) that point diagonally forward and cross in a pinching action, in contrast to the Mygalomorphae (tarantulas and their cl ...
), implicitly adopting the phylogeny shown below. Pocock's approach was criticized by other arachnologists. Thus in 1923, Petrunkevitch rejected grouping mygalomorphs and araneomorphs into Opisthothelae, treating Liphistiomorphae (i.e. Mesothelae), Mygalomorphae and Arachnomorphae (Araneomorphae) as three separate groups. Others, such as Bristowe in 1933, put Liphistiomorphae and Mygalomorphae into one group, called Orthognatha, with Araneomorphae as Labidognatha: In 1976, Platnick and Gertsch argued for a return to Pocock's classification, drawing on morphological evidence. Subsequent phylogenetic studies based on molecular data have vindicated this view. The accepted classification of spiders is now: Order
Araneae Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species dive ...
(spiders) :Suborder Mesothelae Pocock, 1892 :Suborder
Opisthothelae The Opisthothelae are spiders within the order Araneae, consisting of the Mygalomorphae and the Araneomorphae, but excluding the Mesothelae. The Opisthothelae are sometimes presented as an unranked clade and sometimes as a suborder of the Arane ...
Pocock, 1892 ::Infraorder
Mygalomorphae The Mygalomorphae, or mygalomorphs, are an infraorder of spiders, and comprise one of three major groups of living spiders with over 3000 species, found on all continents except Antarctica. Many members are known as trapdoor spiders due to the ...
Pocock, 1892 ::Infraorder
Araneomorphae The Araneomorphae (also called the Labidognatha) are an infraorder of spiders. They are distinguishable by chelicerae (fangs) that point diagonally forward and cross in a pinching action, in contrast to the Mygalomorphae (tarantulas and their cl ...
Smith, 1902 (
syn. The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linnae ...
Arachnomorphae Pocock, 1892)


Distribution

Liphistiinae spiders are distributed in
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 ...
,
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
, the
Malayan peninsula The Malay Peninsula (Malay: ''Semenanjung Tanah Melayu'') is a peninsula in Mainland Southeast Asia. The landmass runs approximately north–south, and at its terminus, it is the southernmost point of the Asian continental mainland. The area ...
, and
Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
. Heptathelinae are found in
Vietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
, the Eastern provinces of China, and Southern Japan.


Fossils

A number of families and genera of fossil
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s have been assigned to the Mesothelae, particularly by
Alexander Petrunkevitch Alexander Ivanovitch Petrunkevitch (Russian: Александр Иванович Петрункевич, December 22, 1875 in Plysky near Kyiv, now Ukraine – March 9, 1964 in New Haven) was an eminent Russian arachnologist of his time. From ...
. However, Paul A. Selden has shown that most only have "the general appearance of spiders", with segmented abdomens (
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 a ...
e), but no definite spinnerets. These families include: * †
Arthrolycosidae Arthrolycosidae is an extinct family of arachnids, possibly spiders. Fossils placed in the family were found in the Carboniferous to Permian, . They were considered by Alexander Petrunkevitch to be "mesotheles", i.e. placed in the spider suborde ...
Frič, 1904 * †
Arthromygalidae The Arthromygalidae are an extinct family of arachnids, possibly spiders. Fossils placed in the family were all found in the Carboniferous, . They were considered by Alexander Petrunkevitch to be "mesotheles", i.e. placed in the spider suborder M ...
Petrunkevitch, 1923 * † Pyritaraneidae Petrunkevitch, 1953 * †''
Palaeothele ''Palaeothele'' is an extinct genus of mesothele spiders, with only one known species ''Palaeothele montceauensis''. Two fossils were found at Montceau-les-Mines, France, in ironstone concretion deposits of Late Carboniferous ( Stephanian) age, ...
'' Selden, 2000 (unplaced in a family) Between 2015 and 2019 six genera of Mesothele spider in four families were described from Late Cretaceous (
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 ...
) aged
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 ...
in
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 ...
. '' Cretaceothele'' (Cretaceothelidae) '' Burmathele''J. Wunderlich. 2017. New and rare fossil spiders (Araneae) in mid Cretaceous amber from Myanmar (Burma), including the description of new extinct families of the suborders Mesothelae and Opisthothelae, as well as notes on the taxonomy, the evolution and the biogeography of the Mesothelae. ''Ten Papers on Fossil and Extant Spiders (Araneae). Beiträge zur Araneologie'' 10:72-279 (Burmathelidae), '' Parvithele, Pulvillothele'' (Parvithelidae) '' Intermesothele'' and '' Eomesothele'' (Eomesothelidae)J. Wunderlich. 2019. What is a spider?. ''Beiträge zur Araneologie'' 12:1-32


References

{{Authority control Carboniferous arthropods Arthropod suborders