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''Parioscorpio'' is an extinct genus of
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 ...
containing the species ''P. venator'' known from 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 Paleozo ...
-aged
Waukesha Biota The Waukesha Biota (also known as Waukesha Lagerstätte, Brandon Bridge Lagerstätte, or Brandon Bridge fauna) refers to a Konservat-Lagerstätte ( a fossil site that preserves soft bodied remains) of Early Silurian (Telychian to Sheinwoodian) a ...
of the
Brandon Bridge Formation The Brandon Bridge Formation is a geologic formation in Wisconsin. It preserves fossils dating back to the Silurian period, including those of graptolites, conodonts and ''Parioscorpio.'' See also * List of fossiliferous stratigraphic units ...
near Waukesha,
Wisconsin Wisconsin () is a state in the upper Midwestern United States. Wisconsin is the 25th-largest state by total area and the 20th-most populous. It is bordered by Minnesota to the west, Iowa to the southwest, Illinois to the south, Lake M ...
.


Taxonomy

The fossils were originally discovered in 1985, tentatively identified as a
branchiopod Branchiopoda is a class of crustaceans. It comprises fairy shrimp, clam shrimp, Diplostraca (or Cladocera), Notostraca and the Devonian ''Lepidocaris''. They are mostly small, freshwater animals that feed on plankton and detritus. Description M ...
or
remipede Remipedia is a class of blind crustaceans found in coastal aquifers which contain saline groundwater, with populations identified in almost every ocean basin so far explored, including in Australia, the Caribbean Sea, and the Atlantic Ocean. The ...
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
but were neglected for decades. In 2016, some of the fossils now assigned to ''Parioscorpio'' were given the name ''Latromirus'' and were assigned to an extinct group of early
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ' ...
arthropods known as
cheloniellid Cheloniellida is a taxon (usually referred to as an order) of extinct Paleozoic arthropods. As of 2018,Wendruff, Andrew James, et al. "New cheloniellid arthropod with large raptorial appendages from the Silurian of Wisconsin, USA." BioRxiv (2018) ...
s in a Ph.D dissertation, but the name was never published in a peer-reviewed journal and is therefore not valid in accordance with the
International Code of Zoological Nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the ...
. Upon initial publication in 2020, ''Parioscorpio'' was considered the world's oldest and most primitive known
scorpion Scorpions are predatory arachnids of the order Scorpiones. They have eight legs, and are easily recognized by a pair of grasping pincers and a narrow, segmented tail, often carried in a characteristic forward curve over the back and always end ...
, older than ''
Dolichophonus ''Dolichophonus'' is an extinct genus of scorpion known from the Silurian aged Gutterford Burn Eurypterid bed, in the Pentland Hills in Scotland. It is currently considered the world's oldest known scorpion, as the older '' Parioscorpio'' name ...
'' from Scotland by several million years. In 2021, the fossils were reanalysed, and ''Parioscorpio'' was found not to be a scorpion, but an arthropod of uncertain placement, outside of
Mandibulata Mandibulata, termed "mandibulates", is a clade of arthropods that comprises the extant subphyla Myriapoda (millipedes and others), Crustacea and Hexapoda (insects and others). Mandibulata is currently believed to be the sister group of the clade ...
,
Chelicerata The subphylum Chelicerata (from New Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. It contains the sea spiders, horseshoe crabs, and arachnids (including Opiliones, harvestmen, scorpions, spiders, Solifugae, solif ...
and all other groups of extinct arthropods (e.g.
Megacheira Megacheira ("great hands") is an extinct class of predatory arthropods that possessed a pair of great appendages, hence the class name as well as the common name "great appendage arthropods". Their taxonomic position is controversial, with stud ...
,
Fuxianhuiida Fuxianhuiida is an extinct clade of arthropods from the Cambrian of China. All currently known species are from Cambrian Series 2 aged deposits in Yunnan Province, including the Chengjiang biota. They are generally considered to be close to the ...
,
Artiopoda The Artiopoda is a grouping of extinct arthropods that includes trilobites and their close relatives. It was erected by Hou and Bergström in 1997 to encompass a wide diversity of arthropods that would traditionally have been assigned to the Trilo ...
and so on). In 2021 another paper stated that ''Parioscorpio venator'', including the fossils previously called ''Latromirus'', might be a
cheloniellid Cheloniellida is a taxon (usually referred to as an order) of extinct Paleozoic arthropods. As of 2018,Wendruff, Andrew James, et al. "New cheloniellid arthropod with large raptorial appendages from the Silurian of Wisconsin, USA." BioRxiv (2018) ...
. If this is correct, it means that ''P. venator'' is related to
trilobites Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the At ...
, nektaspids, aglaspidids, xenopods, and xandarellids. However in 2022, its affinity as cheloniellid is questioned, and firmly rejected from that clade. Currently the most resolved tree in the paper considered ''P. venator'' as an enigmatic
stem-group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
arthropod. In 2022 a study was published describing ''
Acheronauta stimulapis ''Acheronauta'' is a genus of extinct vermiform arthropod that lived in the early Silurian (Telychian-Sheinwoodian stages) Waukesha biota fossil site in southeast Wisconsin. This arthropod was first discovered alongside the biota in 1985, but wa ...
'', a new species of possible mandibulate arthropod from the biota. While coding the phylogenetic trees for this arthropod, the authors of the paper also included ''Parioscorpio'', and all of the trees preformed presented this creature as a basal taxon of arthropod that sat in between the groups artipoda and mandibulata. This discovery actually is consistent with the rejection of ''P. venator'' as a cheloniellid.


Morphology

The animal is around long. It is characterized by a trapezoidal head with a pair of eyes located antero-medially, a pair of enlarged raptorial appendages (previously thought to be scorpion's clawed
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), as well as another pair of small appendages. Central to the head was a mouth-covering
hypostome In zoology, the hypostome can refer to structures in distinct animal groups: *Hypostome (trilobite), the ventral mouthpart plate in trilobites *Hypostome (tick) The hypostome (also called the maxilla, radula, or labium) is a calcified harpoo ...
and a pair of muscular blocks articulated to the raptorial appendages. The trunk is composed of 14 segments, each associated with a pair of thin pleurae (lateral extension of
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 ...
) and appendages. The first segment is covered by the head while the posterior segments may have lateral spines. The anterior 12 pairs of trunk appendages are multiramus (each composed of 4 bundles of setae and a segmented endopod) while the last two pairs are simple fan-like structures. The trunk ends with 3 spines.


Paleoecology

''Parioscorpio'' may had been a marine or brackish water
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
, using an ambush prey-capture method similar to extant waterbugs (
Nepomorpha Nepomorpha is an infraorder of insects in the "true bug" order (Hemiptera). They belong to the "typical" bugs of the suborder Heteroptera. Due to their aquatic habits, these animals are known as true water bugs. They occur all over the world o ...
). It would have lived alongside many other bizarre organisms like the
Conodont Conodonts (Greek ''kōnos'', "cone", + ''odont'', "tooth") are an extinct group of agnathan (jawless) vertebrates resembling eels, classified in the class Conodonta. For many years, they were known only from their tooth-like oral elements, which ...
''
Panderodus ''Panderodus'' Is an extinct genus of jawless fish belonging to the order Conodonta. This genus had a long temporal range, surviving from the middle Ordovician to late Devonian. In 2021, extremely rare body fossils of ''Panderodus'' from the Wauke ...
,'' the enigmatic Butterfly Animal, the Thylacocephalan '' Thylacares,'' early synziphosurans, and Trilobites.


References

{{Taxonbar, from=Q83291300 Silurian life of North America Telychian first appearances Fossil taxa described in 2020 Prehistoric arthropod genera Controversial taxa Silurian arthropods of North America