Pentecopterus Temporal range: Darriwilian, | |
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Reconstruction of Pentecopterus decorahensis, showing a dorsal view (a), a genital operculum (b), a ventral view of carapace (c) and a ventral view of prosoma (d). | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Subphylum: | Chelicerata |
Order: | † Eurypterida |
Family: | † Megalograptidae |
Genus: | † Pentecopterus Lamsdell et al., 2015 |
Type species | |
†Pentecopterus decorahensis Lamsdell et al., 2015 |
Pentecopterus is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils have been registered from the Darriwilian age of the Middle Ordovician period, as early as 467.3 million years ago. The genus contains only one species, P. decorahensis, that is the oldest known eurypterid, surpassing other Ordovician eurypterids, such as Brachypterus , in age by almost 9 million years. The generic name derives from the penteconter, a warship from ancient Greece, and the suffix -pterus, which means "wing" and is often used in other genus of eurypterids. The specific name refers to Decorah, Iowa, where Pentecopterus was discovered.
The genus is classified as part of the Megalograptidae family of eurypterids, a family differentiated from other eurypterids by the possession of two or more pairs of spines per podomere on prosomal appendage IV, a reduction of almost all spines and the large exoskeletons with ovate to triangular scales. [1] It is estimated that Pentecopterus had a length of 1.7 metres (5 ft 7 in), turning it into one of the largest eurypterids or arthropods ever discovered. However, Pentecopterus was overtaken by other eurypterids such as Jaekelopterus rhenaniae , the largest known arthropod with 2.5 metres (8 ft 2 in).
A replica of the eurypterid was created for the National Geographic's " The Strange Truth " program. The project had the help of James Lamsdell, one of the paleontologists who contributed to the description of Pentecopterus, who provided information and two illustrations. When it was finished, it was taken to the impact site where the original fossils were discovered. [2]
Pentecopterus is among the largest known arthropods, with the largest specimens having an estimated length of 1.7 metres (5 ft 7 in). With this size, it exceeds all other genera of the Megalograptidae. While fragmentary tergites have been suggested to indicate a length of 2 metres (6 ft 7 in) for Megalograptus shideleri , they actually indicate a length of at most 56 cm (22 in). [3] [4] Most Pentecopterus specimens would have had a total length of 75–100 cm (30–39 in). [3]
The large amount of fragmentary specimens recovered of Pentecopterus, including juveniles and exuviae specimens, have allowed an almost complete description of the external morphology. Pentecopterus is diagnosed as a megalograptid retaining a single pair of spines on the third podomere of the third prosomal appendages, a short appendage V with a serrated distal margin of podomeres; prosomal ventral plates widening anteriorly, posterolateral pretelson lacking expansion and xiphos-like shaped telson, with a margin laterally ornamented with scales. The prosomal ventral plate is of Erieopterus -type, that is, it consists of a single plate that covers the anterior and lateral portion of the ventral carapace. The appendages are attached to the soft ventral integument of the plate. The general outline of the carapace was quadrate with an elongate trapezoidal outline. [3]
Most of the fossils of the Winneshiek Shale of Decorah, where the fossils of Pentecopterus have been found, were recovered in 2010 from the upper 4 m section of the aforementioned site in Iowa. Other samples were collected from blocks eroded during flooding, which are assumed to have been sourced from the uppermost 2–3 m. The total number of fossils found is over 5,000, of which around 6.6% belong to eurypterids. Among the material eurypterid is the holotype of Pentecopterus, SUI 139941, which consists of a prosomal ventral plate and proximal podomeres of prosomal appendage II, in addition to multiple paratypes and additional material, all housed in the University of Iowa Paleontology Repository. [3]
The genus was named by James C. Lamsdell, Derek E. G. Briggs, Huaibao P. Liu, Brian J. Witzke and Robert M. McKay in 2015 and the type species is Pentecopterus decorahensis. It is the only eurypterid discovered in Iowa other than Mycterops whitei , [5] a species whose assignment to Mycterops is doubtful. The genus is named after the penteconter, an early galley (ship) from ancient Greece and one of the first true warships, due to some similarities in shape and that the taxon represents an early predator. The suffix -pterus means "wing" and is commonly applied to eurypterid genera. The specific name decorahensis refers to Decorah, Iowa, where the fossils were discovered. [3]
Pentecopterus is classified within the family Megalograptidae in the monotypic superfamily Megalograptoidea. Pentecopterus shares with the other megalograptids a series of characteristics such as the randomly-oriented armature on the swollen podomeres of appendage IV and the narrow gnathobase bearing multiple rows of small teeth on the coxa of appendage V. It also shares with Megalograptus the rows of enlarged scales running down the center of the opisthosomal tergites, however, these differ from those of Megalograptus because they are not situated on pronounced ridges. It is separated from the other megalograptids by the presence of only one pair of spines on the third podomere of appendage III, a single terminal spine on each prosomal appendage, and the absence of dense cuticular ornamentation. Pentecopterus also lacks the cercal blades that occur in Megalograptus, where they have been interpreted as functioning as a biological rudder, like the pterygotid telson. This suggests that Pentecopterus was less able to swim than Megalograptus. Further, Pentecopterus has some features unique among the eurypterids, notably the shape of its carapace and the unusual shape of the sixth podomere of appendage VI. Pentecopterus also possesses lateral scales on the telson, something otherwise only seen in pterygotid eurypterids. This feature likely arose through convergent evolution. [3]
The cladogram below is simplified from a study by Lamsdell et al. (2015), [3] showing the phylogenetic positions of the genera within Megalograptidae.
Megalograptidae |
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Pentecopterus is known from Middle Ordovician deposits in the Winneshiek Shale of Iowa dated to the Darriwilian (467.3 mya). The Winneshiek Shale is an 18–27 m thick greenish brown to dark grey laminated sandy shale that crops out only in one locality which is mostly submerged by the Upper Iowa River near Decorah. Bore hole data indicate that the total thickness of the Winneshiek Shale is about 18 m at the outcrop locality, but only the upper 4 m was systematically collected during the excavation. In these 4 m, most of the Pentecopterus fossils were collected. This shale is confined to a circular basin about 5.6 km in diameter in the Decorah area that probably originated from a meteorite impact. Palaeogeographic and paleoenvironmental studies suggest that this crater was located in marginal to nearshore marine conditions with little oxygen, possibly in a brackish water environment within the tropical southern Laurentia. The Winneshiek fauna is very different from the shelly marine fossil fauna, which indicates that this environment was inhospitable to the typical marine taxa. This fauna was composed of conodonts, arthropods, possible jawless fishes and algae. [3]
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.
Megalograptus is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Megalograptus have been recovered in deposits of Katian 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 "great writing" and originates from the mistaken original belief that Megalograptus was a type of graptolite, often given names ending with -graptus.
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.
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.
Onychopterella is a genus of predatory eurypterid, an extinct group of aquatic arthropods. Fossils of Onychopterella have been discovered in deposits from the Late Ordovician to the Late Silurian. The genus contains three species: O. kokomoensis, the type species, from the Early Pridoli epoch of Indiana; O. pumilus, from the Early Llandovery epoch of Illinois, both from the United States; and O. augusti, from the Late Hirnantian to Early Rhuddanian stages of South Africa.
Holmipterus is a problematic genus of eurypterid, an extinct group of aquatic arthropods. The type and only species of Holmipterus, H. suecicus, is known from deposits of Middle Silurian age in the Sweden. The generic name honours Gerhard Holm, a renowned Swedish palaeontologist specialising in arthropods and crustaceans, and the species name suecicus is Latin for 'Swedish'.
Eocarcinosoma is a genus of eurypterid, an extinct group of aquatic arthropods. The type and only species of Eocarcinosoma, E. batrachophthalmus, is known from deposits of Late Ordovician age in the United States. The generic name is derived from the related genus Carcinosoma, and the Greek eós meaning 'dawn', referring to the earlier age of the genus compared to other carcinosomatid eurypterids.
Echinognathus is a genus of eurypterid, an extinct group of aquatic arthropods. The type and only species of Echinognathus, E. clevelandi, is known from deposits of Late Ordovician age in the United States. The generic name is derived from the Neo-Latin echino- ("spiny") and the Greek gnáthos ("jaw"), in reference to a spiny endognathary appendage part of the fossil type material.
Rhinocarcinosoma is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Rhinocarcinosoma have been discovered in deposits ranging of Late Silurian age in the United States, Canada and Vietnam. The genus contains three species, the American R. cicerops and R. vaningeni and the Vietnamese R. dosonensis. The generic name is derived from the related genus Carcinosoma, and the Greek ῥινός, referring to the unusual shovel-shaped protrusion on the front of the carapace of Rhinocarcinosoma, its most distinctive feature.
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 are a family of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions".
The Decorah crater, also called the Decorah impact structure, is a possible impact crater located on the east side of the city of Decorah in Iowa, United States. It is thought to have been caused by a meteor about 200 metres (660 ft) wide which struck during the Middle Ordovician Period, circa 470 million years ago.
Carcinosomatoidea is an extinct superfamily of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". It is one of the superfamilies classified as part of the suborder Eurypterina.
Hughmilleriidae is a family of eurypterids, an extinct group of aquatic arthropods. The hughmilleriids were the most basal members of the superfamily Pterygotioidea, in contrast with the more derived families Pterygotidae and Slimonidae. Despite their classification as pterygotioids, the hughmilleriids possessed several characteristics shared with other eurypterid groups, such as the lanceolate telson.
Herefordopterus is a genus of eurypterid, an extinct group of aquatic arthropods. Herefordopterus is classified as part of the family Hughmilleriidae, a basal family in the highly derived Pterygotioidea superfamily of eurypterids. Fossils of the single and type species, H. banksii, have been discovered in deposits of Silurian age in Herefordshire and Shropshire, England. The genus is named after Herefordshire, where most of the Herefordopterus fossils have been found. The specific epithet honors Richard Banks, who found several well-preserved specimens, including the first Herefordopterus fossils.
Wiedopterus is a genus of eurypterid, an extinct group of aquatic arthropods. The type and only species of Wiedopterus, W. noctua, is known from deposits of Early Devonian age in Germany. The generic name derives from the Wied river, which runs near the site of the initial discovery, and the species name, noctua, derives from Latin noctua (owl) which refers to the superficial resemblance of the carapace to an owl.
Winneshiekia is an extinct genus of dekatriatan, a clade of chelicerate arthropods. Fossils of the single and type species, W. youngae, have been discovered in deposits of the Middle Ordovician period in Iowa, in the United States. The name of the genus is derived from the Winneshiek Shale, the formation in which it was discovered. The species name youngae honors Jean N. Young, an American geologist who contributed greatly to the discovery of the Winneshiek Shale.
Camanchia is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Camanchia was regarded as part of the clade Prosomapoda. Fossils of the single and type species, C. grovensis, have been discovered in deposits of the Silurian period in Iowa, in the United States. Alongside Venustulus, Camanchia is one of the only Silurian synziphosurine with fossil showing evidence of appendages.
The Winneshiek Shale is a Middle Ordovician (Darriwilian-age) geological formation in Iowa. The formation is restricted to the Decorah crater, an impact crater near Decorah, Iowa. Despite only being discovered in 2005, the Winneshiek Shale is already renowned for the exceptional preservation of its fossils. The shale preserves a unique ecosystem, the Winneshiek biota, which is among the most remarkable Ordovician lagerstätten in the United States. Fossils include the oldest known eurypterid, Pentecopterus, as well as giant conodonts such as Iowagnathus and Archeognathus.
Terropterus is a genus of eurypterid, an extinct group of aquatic arthropods. The type and only species of Terropterus, T. xiushanensis, is known from deposits of Early Silurian age in China.