Arthropleuridea
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Arthropleuridea is an extinct subclass of
myriapod Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. The fossil record of myriapods reaches back into the late Silurian, ...
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 that flourished during 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 ''Carboniferou ...
period, having first arose during the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleoz ...
, and perishing in the
Early Permian 01 or '01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * 01'' (Richard Müller album), 2001 * ''01'' (Son of Dave album), 2000 * ''01'' (Urban Zakapa album), 2011 * ''O1'' (Hiroyuki Sawan ...
. Members are characterized by possessing diplosegement (fused "double segments", as in modern-day
millipedes Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a resu ...
) paranotal tergal lobes separated from the body axis by a suture, and by sclerotized plates buttressing the leg insertions. Despite their unique features, recent
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
research suggests Arthropleuridea be included among millipedes in the class Diplopoda. The subclass contains three recognized
orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
, each with a single
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
.


Paleobiology

Arthropleurids lived in the moist
coal Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen. Coal is formed when ...
swamps that were common at the time and may have
burrow An Eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of locomotion. Burrows provide a form of s ...
ed in the undergrowth. They were either
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
s or
detritivore Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrate ...
s. Besides their size, their most distinguishing features were their legs with eight segments (as many as 30 pairs) and extremely tough
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 ...
s. There is no evidence of spiracles, so the animals must have used
lung The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of ...
s or
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they ar ...
s for
respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
. Most arthropleurideans are thought to have been terrestrial, although, without any known respiratory structure, terrestriality is assumed only by analogy to modern arthropods. Early forms, however, including '' Eoarthropleura'' (order Eoarthropleurida), appear to have been aquatic. For this reason, some question Arthropleuridea's inclusion among millipedes because no modern aquatic
myriapods Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. The fossil record of myriapods reaches back into the late Silurian, ...
are known.


Distribution and size

Arthropleuridea is most famous for ''
Arthropleura ''Arthropleura'' () is a genus of extinct millipede arthropods that lived in what is now North America and Europe around 345 to 290 million years ago, from the Viséan stage of the lower Carboniferous Period to the Sakmarian stage of the lower ...
'' (order Arthropleurida). Tracks from ''Arthropleura'' up to 50 cm wide have been found at
Joggins, Nova Scotia Joggins is a rural community located in western Cumberland County, Nova Scotia, Canada. On July 7, 2008 a 15-km length of the coast constituting the Joggins Fossil Cliffs was officially inscribed on the World Heritage List.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 ever to have lived. The lack of large terrestrial vertebrate predators and the highly
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements ...
ic atmosphere at that time probably enabled them to grow so large. It was previously believed that ''Arthropleura'' became
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
as the coal swamps dried out. However, many fossils have been discovered after the Carboniferous rainforest collapse. A more recent proposal is that predation by
tetrapods Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct therapsi ...
and the
aridification Aridification is the process of a region becoming increasingly arid, or dry. It refers to long term change, rather than seasonal variation. It is often measured as the reduction of average soil moisture content. It can be caused by reduced precip ...
of the
equator The equator is a circle of latitude, about in circumference, that divides Earth into the Northern and Southern hemispheres. It is an imaginary line located at 0 degrees latitude, halfway between the North and South poles. The term can also ...
caused it to become extinct. ''Arthropleura'' has been found from the Upper
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 ''Carboniferou ...
of
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
and
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
. '' Eoarthropleura'' has been found from the Upper Silurian through the Upper
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
of
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
and
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
. ''
Microdecemplex ''Microdecemplex rolfei'' is the only known species of the extinct, small-bodied millipede order Microdecemplicida, a member of the extinct subclass Arthropleuridea. Fossils, measuring less than 10 mm in length, are known from the Panther ...
'', of the order Microdecemplicida, was dwarfed by the other Arthropleurideans, at just a few millimeters long. The genus is known from the Middle through Upper
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
of
New York New York most commonly refers to: * New York City, the most populous city in the United States, located in the state of New York * New York (state), a state in the northeastern United States New York may also refer to: Film and television * '' ...
state, USA.


Classification and placement

After several decades of uncertainty, Arthropleuridea was placed within the Diplopoda in the year 2000. However, there is still controversy regarding the relationships of the three orders to living millipede groups. Some authors place Arthropleuridea within the
Chilognatha Chilognatha is a subclass of the class Diplopoda, which includes the vast majority of extant millipedes, about 12,000 species. Taxonomy The classification of Chilognatha presented below is based on Shear, 2011, and Shear & Edgecombe, 2010 (exti ...
, as a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to all living Chilognathan millipedes (
Pentazonia Pentazonia is a taxonomic infraclass of millipedes containing the pill-millipedes (Oniscomorpha) which can roll into a ball and the order Glomeridesmida which cannot. Defining traits (apomorphies) include divided sternites, a labrum with single ...
+
Helminthomorpha Chilognatha is a subclass of the class Diplopoda, which includes the vast majority of extant millipedes, about 12,000 species. Taxonomy The classification of Chilognatha presented below is based on Shear, 2011, and Shear & Edgecombe, 2010 (exti ...
). An alternate hypothesis breaks up the subclass: placing the orders Arthropleurida and Eoarthropleurida within the basal Penicillata (as sister to the living
Polyxenida Polyxenida is an order of millipedes readily distinguished by a unique body plan consisting of a soft, non-calcified body ornamented with tufts of bristles – traits that have inspired the common names "bristly millipedes" or "pincushion millipe ...
), and leaving only Microdecemplicida as a sister group to the living Chilognatha. Under this hypothesis, Arthropleuridea would be
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
.


See also

*
Archipolypoda Archipolypoda is an extinct group of millipedes known from fossils in Europe and North America and containing the earliest known land animals. The Archipolypoda was erected by Scudder (1882) but redefined in 2005 with the description of sever ...
, another extinct group of millipedes including the earliest known air-breathing animal * Euthycarcinoidea, a group of enigmatic arthropods that may be ancestral to myriapods *
Colonization of land The history of life on Earth traces the processes by which living and fossil organisms evolved, from the earliest emergence of life to present day. Earth formed about 4.5 billion years ago (abbreviated as ''Ga'', for ''gigaannum'') and evide ...
, major evolutionary stages leading to terrestrial organisms


References

{{Taxonbar, from=Q4797674 Millipede taxonomy Arthropod subclasses Prehistoric arthropod taxa Carboniferous myriapods Devonian myriapods Silurian myriapods Pridoli first appearances