Eurypterina

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Eurypterina
Temporal range: Middle Ordovician - Early Permian, 467.3–283.5  Ma
Bassipterus virginicus swimming leg.png
The defining characteristic of eurypterine eurypterids is the transformation of the sixth pair of legs into swimming paddles. Reconstructed leg of Bassipterus .
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Clade: Sclerophorata
Order: Eurypterida
Suborder: Eurypterina
Burmeister, 1843
Type species
Eurypterus remipes
De Kay, 1825
Subgroups
Incertae sedis
Synonyms
  • Pterygotina Caster & Kjellesvig-Waering, 1964

Eurypterina is one of two suborders of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". Eurypterine eurypterids are sometimes informally known as "swimming eurypterids". [1] They are known from fossil deposits worldwide, though primarily in North America and Europe.

Contents

Seventy-five percent of eurypterid species are eurypterines; this represents 99% of specimens. [2] The superfamily Pterygotioidea is the most species-rich clade, with 56 species, followed by the Adelophthalmoidea with 43 species; as sister taxa, they comprise the most derived eurypterines. Pterygotioidea includes the pterygotids, which are the only eurypterids known to have a cosmopolitan distribution. [3]

Though more numerous both in specimens and taxa, the eurypterines have the shorter temporal range of the two eurypterid suborders. They first appeared around the same time as the Stylonurina in the Middle Ordovician. The suborder faced a slow extinction during the Middle and Late Devonian, possibly tied to the emergence of jawed vertebrates. [4] Every Eurypterine genus and lineage went extinct before the Carboniferous save for Adelophthalmus which would go extinct in the Early Permian, millions of years before the Permian-Triassic extinction event that ended the stylonurines. [5]

Description

The Stylonurina and Eurypterina are most easily distinguished by the morphology of the posteriormost prosomal appendage. In the Stylonurina, this appendage takes the form of a long and slender walking leg, lacking a modified spine (termed podomere 7a). In the Eurypterina, the leg is most usually modified and broadened into a swimming paddle and always includes a podomere 7a. [3]

Swimming eurypterines represent the absolute majority of both known eurypterid species and known specimens, though the morphology of the walking stylonurines is almost as diverse in appearance, and the fossil record of the eurypterines may therefore simply be more complete than that of the stylonurines, possibly due to varying habitat preferences. [3]

Paleobiogeography

Life restoration of Eurypterus. Eurypterus is by far the most well-studied and well-known eurypterid and its fossil specimens probably represent more than 90 % of all known eurypterid specimens. Eurypterus Paleoart.jpg
Life restoration of Eurypterus . Eurypterus is by far the most well-studied and well-known eurypterid and its fossil specimens probably represent more than 90 % of all known eurypterid specimens.

The most basal eurypterines with swimming legs, the genus Onychopterella , are known from the east coast of Gondwana close to the equator (a region that today is South Africa) from the Late Ordovician. It is not known whether or not the swimming forms originated here or not, but it is speculated that they migrated from Laurentia, since most stylonurines and basal swimming forms are predominantly known from Laurentia and Gondwana otherwise completely lacks basal swimming forms. [3]

The megalograptoids were likely the first major successful group of eurypterids, evidenced by a Late Ordovician radiation. All known members of the Megalograptoidea are from the Middle to Late Ordovician of Laurentia, though potential records from the Middle Silurian of Baltica are known in the form of the genus Holmipterus suecicus (though its classification as a megalograptoid is questionable). [3]

Eurypteroids are known from Laurentia and Baltica, with one known species from Avalonia. Eurypterus and other eurypteroids appear to have been unable to spread beyond Laurussian waters. The genus Eurypterus in particular dominated many Silurian eurypterid faunas of Laurentia. Despite its abundance, it appears to not have originated in Laurentia, the earliest records of the genus are from Baltica and Eurypterus was thus likely an invasive genus in Laurentia, albeit one that managed to adapt well to the new habitats. [3]

The majority of carcinosomatoid taxa are also known from Laurentia, Baltica and Avalonia. Isolated and fragmentary fossils from the Late Silurian of Vietnam and the Czech Republic show that the terranes of Annamia and Perunica were within the geographical range of the carcinosomatoids. Only a few basal carcinosomatoids (e.g. Carcinosoma and Paracarcinosoma ) have been found in deeper waters whilst the more derived forms, such as Mixopterus and Lanarkopterus have not. Basal carcinosomatoids (Carcinosomatidae) are likely responsible for the fossil remains in Vietnam and the Czech Republic and may have had a distribution similar to the cosmopolitan distribution of the pterygotoids, though were not as common nor as successful. [3]

Adelophthalmoids were the longest lasting clade of eurypterines, becoming extinct in the Middle Permian, this is in part due to the survival of Adelophthalmus beyond the Middle Devonian. The earliest records of the genus are from the Early Devonian of western Germany, but following the amalgamation of Pangaea during the Carboniferous and Permian, the genus gained an almost cosmopolitan distribution. The basalmost species in the entire clade are from Baltica and most of the evolution within the basal members took place in Laurussia. By the Devonian, representatives were found in both Siberia and Australia long before the formation of Pangaea. [3]

Although the Pterygotoidea only existed for a period of about 40 million years during a time when most continents were widely separated, the clade is the eurypterid clade with the most cosmopolitan distribution. Like other eurypterines, they are most common in Laurentia, Baltica and Avalonia, but are also found commonly in other paleocontinents. Fossil remains have been recovered from Australia, Libya, Algeria, Morocco, Florida, Saudi Arabia, Iberia, South America, vast swaths of Gondwana, Bohemia and Siberia. The earliest pterygotoids are from the latest Llandovery of Scotland, Laurentia and South China and this mobility makes it difficult to pinpoint the geographical origin of the clade, though it is speculated to have been close to or in Laurentia like the Adelophthalmoidea. [3]

Systematics and relationships

Fossil of Mixopterus, a mixopterid. Mixopterus.jpg
Fossil of Mixopterus , a mixopterid.

Eurypterina contains eight superfamilies - Onychopterelloidea, Moselopteroidea, Megalograptoidea, Eurypteroidea, Carcinosomatoidea, Waeringopteroidea, Adelophthalmoidea and Pterygotioidea. The relationships between them remain somewhat unclear, the Megalograptoidea is thought to be relatively primitive (between Onychopterella and the Eurypteroidea) because they lack a synapomorphy of all more derived swimming forms; the modified distal margin of the sixth podomere of the swimming leg. This position is not necessarily true, since the sixth podomere in the swimming leg resembles the reduced podomere found in the Mixopteridae, and they might instead belong between the Eurypteroidea and Carcinosomatoidea. [3]

In contrast to the Megalograptoidea, the Eurypteroidea is a rather well-known clade that contains around 90% of all known eurypterid specimens. They were closely related, supported by numerous similarities, to the Carcinosomatoidea. The Carcinosomatoidea have a poorly resolved internal phylogeny, though can be easily recognised by scorpion-like appearance and heavily spinose appendages. [3]

Pterygotioidea and Adelophthalmoidea are the two most derived clades as well as the most taxonomically diverse ones. Adelophthalmoidea contains 43 species, whereas Pterygotioidea contains 56. [3] The superfamilies classified as part of Eurypterina contain the following families:

Fossil of Jaekelopterus, a pterygotioid. At a length of 2.5 meters, it is one of the largest arthropods to have ever lived. Jaekelopterus rhenaniae.jpg
Fossil of Jaekelopterus , a pterygotioid. At a length of 2.5 meters, it is one of the largest arthropods to have ever lived.

Suborder Eurypterina Burmeister, 1843

Phylogeny

Eurypterines are characterised by the transformation of the posteriormost prosomal appendage into a swimming paddle, one of the main features used to distinguish them from the stylonurines. The cladogram presented below, simplified from a study by Tetlie, [3] showcases the phylogenetic relationships of the Eurypterina based on this adaptation, and the enlargement of the chelicerae, which characterises the family Pterygotidae, to be used for active prey capture.

Eurypterida

Stylonurina

Eurypterina

Megalograptoidea

Eurypteroidea

Carcinosomatoidea

Waeringopteroidea

Adelophthalmoidea

Pterygotioidea

Hughmilleria

Herefordopterus

Slimonia

Pterygotidae

See also

Related Research Articles

Eurypterid Order of arthropods (fossil)

Eurypterids, often informally called sea scorpions, are a group of extinct arthropods that form the order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period 467.3 million years ago. The group is likely to have appeared first either during the Early Ordovician or Late Cambrian period. With approximately 250 species, the Eurypterida is the most diverse Paleozoic chelicerate order. Following their appearance during the Ordovician, eurypterids became major components of marine faunas during the Silurian, from which the majority of eurypterid species have been described. The Silurian genus Eurypterus accounts for more than 90% of all known eurypterid specimens. Though the group continued to diversify during the subsequent Devonian period, the eurypterids were heavily affected by the Late Devonian extinction event. They declined in numbers and diversity until becoming extinct during the Permian–Triassic extinction event 251.9 million years ago.

<i>Megalograptus</i> Extinct genus of arthropods

Megalograptus is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Megalograptus have been recovered in deposits of Late Ordovician age in North America. The genus contains five species: M. alveolatus, M. ohioensis, M. shideleri, M. welchi and M. williamsae, all based on fossil material found in the United States. Fossils unassigned to any particular species have also been found in Canada. The generic name translates to "big writing" and originates from the mistaken original belief that Megalograptus was a type of graptolite, a group which typically leave writing-like fossil remains.

Stylonurina Extinct suborder of arthropods

Stylonurina is one of two suborders of eurypterids, a group of extinct arthropods commonly known as "sea scorpions". Members of the suborder are collectively and informally known as "stylonurine eurypterids" or "stylonurines". They are known from deposits primarily in Europe and North America, but also in Siberia.

<i>Carcinosoma</i> Extinct genus of arthropods

Carcinosoma is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Carcinosoma are restricted to deposits of late Silurian age. Classified as part of the family Carcinosomatidae, which the genus lends its name to, Carcinosoma contains seven species from North America and Great Britain.

<i>Hughmilleria</i> Genus of extinct arthropods

Hughmilleria is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Hughmilleria have been discovered in deposits of the Silurian age in China and the United States. Classified as part of the basal family Hughmilleriidae, the genus contains three species, H. shawangunk from the eastern United States, H. socialis from Pittsford, New York, and H. wangi from Hunan, China. The genus is named in honor of the Scottish geologist Hugh Miller.

<i>Drepanopterus</i> Extinct genus of sea scorpions

Drepanopterus is an extinct genus of eurypterid and the only member of the family Drepanopteridae within the Mycteropoidea superfamily. There are currently three species assigned to the genus. The genus has historically included more species, with nine species associated with the genus Drepanopterus, however five of these have since been proven to be synonyms of pre-existing species, assigned to their own genera, or found to be based on insubstantial fossil data. The holotype of one species proved to be a lithic clast.

<i>Pittsfordipterus</i> Genus of arthropods (fossil)

Pittsfordipterus is a genus of eurypterid, an extinct group of aquatic arthropods. Pittsfordipterus is classified as part of the family Adelophthalmidae, the only clade in the derived ("advanced") Adelophthalmoidea superfamily of eurypterids. Fossils of the single and type species, P. phelpsae, have been discovered in deposits of Silurian age in Pittsford, New York state. The genus is named after Pittsford, where the two only known specimens have been found.

<i>Tylopterella</i> Extinct genus of arthropods

Tylopterella is a genus of eurypterid, a group of extinct aquatic arthropods. Only one fossil of the single and type species, T. boylei, has been discovered in deposits of the Late Silurian period in Elora, Canada. The name of the genus is composed by the Ancient Greek words τύλη (túlē), meaning "knot", and πτερόν (pteron), meaning "wing". The species name boylei honors David Boyle, who discovered the specimen of Tylopterella.

<i>Parahughmilleria</i> Extinct genus of arthropods

Parahughmilleria is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Parahughmilleria have been discovered in deposits of the Devonian and Silurian age in the United States, Canada, Russia, Germany, Luxembourg and Great Britain, and have been referred to several different species. The first fossils of Parahughmilleria, discovered in the Shawangunk Mountains in 1907, were initially assigned to Eurypterus. It would not be until 54 years later when Parahughmilleria would be described.

Carcinosomatidae Extinct family of arthropods

Carcinosomatidae is a family of eurypterids, an extinct group of aquatic arthropods. They were members of the superfamily Carcinosomatoidea, also named after Carcinosoma. Fossils of carcinosomatids have been found in North America, Europe and Asia, the family possibly having achieved a worldwide distribution, and range in age from the Late Ordovician to the Early Devonian. They were among the most marine eurypterids, known almost entirely from marine environments.

Megalograptidae Family of arthropods (fossil)

Megalograptidae are a family of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions".

Stylonuridae Extinct family of arthropods

Stylonuridae is a family of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". The family is one of two families contained in the superfamily Stylonuroidea, which in turn is one of four superfamilies classified as part of the suborder Stylonurina.

Rhenopteridae Extinct family of arthropods

The Rhenopteridae are a family of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". The family is the only family currently contained in the superfamily Rhenopteroidea, one of four superfamilies classified as part of the suborder Stylonurina.

Pterygotioidea Extinct superfamily of eurypterids

Pterygotioidea is a superfamily of eurypterids, an extinct group of aquatic arthropods. Pterygotioids were the most derived members of the infraorder Diploperculata and the sister group of the adelophthalmoid eurypterids. The group includes the basal and small hughmilleriids, the larger and specialized slimonids and the famous pterygotids which were equipped with robust and powerful cheliceral claws.

Stylonuroidea Extinct superfamily of arthropods

Stylonuroidea is an extinct superfamily of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". It is one of four superfamilies classified as part of the suborder Stylonurina.

Parastylonuridae Extinct family of arthropods

The Parastylonuridae are a family of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". The family is one of two families contained in the superfamily Stylonuroidea, which in turn is one of four superfamilies classified as part of the suborder Stylonurina.

Waeringopteridae Extinct family of arthropods

Waeringopteridae is a family of eurypterids, an extinct group of aquatic arthropods. The Waeringopteridae is the only family classified as part of the superfamily Waeringopteroidea, which in turn is classified within the infraorder Diploperculata in the suborder Eurypterina. The earliest known member of the group, Orcanopterus, has been recovered from deposits of Katian age and the latest known surviving member, Grossopterus, has been recovered from deposits of Siegenian age. The name Waeringopteridae is derived from the type genus Waeringopterus, which is named in honor of eurypterid researcher Erik N. Kjellesvig-Waering.

Adelophthalmidae Family of eurypterids, an extinct group of aquatic arthropods

Adelophthalmidae is a family of eurypterids, an extinct group of aquatic arthropods. Adelophthalmidae is the only family classified as part of the superfamily Adelophthalmoidea, which in turn is classified within the infraorder Diploperculata in the suborder Eurypterina.

Mycteropoidea Extinct superfamily of arthropods

Mycteropoidea is an extinct superfamily of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". It is one of four superfamilies classified as part of the suborder Stylonurina. Mycteropoids have been recovered from Europe, Russia, South America and South Africa. Mycteropoid specimens are often fragmentary, making it difficult to establish relationships between the included taxa. Only two mycteropoid taxa are known from reasonable complete remains, Hibbertopterus scouleri and H. wittebergensis.

<i>Borchgrevinkium</i> Extinct genus of arthropods

Borchgrevinkium is an extinct genus of chelicerate arthropod. A fossil of the single and type species, B. taimyrensis, has been discovered in deposits of the Early Devonian period in the Krasnoyarsk Krai, Siberia, Russia. The name of the genus honors Carsten Borchgrevink, an Anglo-Norwegian explorer who participated in many expeditions to Antarctica. Borchgrevinkium represents a poorly known genus whose affinities are uncertain.

References

  1. Tetlie, O. Erik; Cuggy, Michael B. (2007). "Phylogeny of the basal swimming eurypterids (Chelicerata; Eurypterida; Eurypterina)". Journal of Systematic Palaeontology. 5 (3): 345–356. doi:10.1017/S1477201907002131. ISSN   1477-2019.
  2. Dunlop JA, Penney D, Tetlie OE, Anderson LI (2008). "How many species of fossil arachnids are there?". Journal of Arachnology . 36 (2): 267–272. doi:10.1636/CH07-89.1. ISSN   0161-8202. S2CID   42371883.
  3. 1 2 3 4 5 6 7 8 9 10 11 12 13 Tetlie OE (2007). "Distribution and dispersal history of Eurypterida (Chelicerata)" (PDF). Palaeogeography, Palaeoclimatology, Palaeoecology . 252 (3–4): 557–574. doi:10.1016/j.palaeo.2007.05.011. Archived from the original (PDF) on 2011-07-18.
  4. Lamsdell JC, Braddy SJ (April 2010). "Cope's Rule and Romer's theory: patterns of diversity and gigantism in eurypterids and Palaeozoic vertebrates". Biology Letters. 6 (2): 265–9. doi:10.1098/rsbl.2009.0700. PMC   2865068 . PMID   19828493.
  5. Lamsdell JC, Braddy SJ, Tetlie OE (2010). "The systematics and phylogeny of the Stylonurina (Arthropoda: Chelicerata: Eurypterida)". Journal of Systematic Palaeontology . 8 (1): 49–61. doi:10.1080/14772011003603564. S2CID   85398946.