Ciurcopterus Temporal range: Late Silurian, | |
---|---|
Fossil specimen of C. ventricosus | |
Restoration of C. ventricosus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Subphylum: | Chelicerata |
Order: | † Eurypterida |
Superfamily: | † Pterygotioidea |
Family: | † Pterygotidae |
Genus: | † Ciurcopterus Tetlie & Briggs, 2009 |
Type species | |
†Ciurcopterus ventricosus Kjellesvig-Waering, 1948 | |
Species | |
|
Ciurcopterus is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Ciurcopterus have been discovered in deposits of Late Silurian age in North America. Classified as part of the family Pterygotidae, the genus contains two species, C. sarlei from Pittsford, New York and C. ventricosus from Kokomo, Indiana. [1] The genus is named in honor of Samuel J. Ciurca, Jr., who has contributed significantly to eurypterid research by discovering a large amount of eurypterid specimens, including the four specimens used to describe Ciurcopterus itself. [2]
Ciurcopterus is the most basal (primitive) known member of the Pterygotidae, and combined characteristics of more derived members of the family with features of close relatives of the group, such as Slimonia . Measuring 70 centimetres (28 inches) in length, Ciurcopterus was relatively large though smaller than many of the later members of its family, which would grow to become the largest known arthropods to have ever lived.
Ciurcopterus was a eurypterid of medium size, with C. ventricosus measuring approximately 70 centimetres (28 in) in length and C. sarlei measuring 50 centimetres (20 in). [4] Though this is large relative to most modern day arthropods, Ciurcopterus was dwarfed by many of the members of its family (the Pterygotidae), such as Jaekelopterus rhenaniae at 2.5 metres (8 ft, the largest known arthropod) and Acutiramus bohemicus at 2.1 metres (7 ft). [5]
Ciurcopterus possessed walking legs that were similar to those of Slimonia in bearing distal serrations. The telson (the posteriormost segment of its body) was wide and possessed dorsal median carinae. The type A genital appendage (one of the morphs of eurypterid genital appendages, equipped with clasping organs) was undivided and the pretelson (the segment immediately preceding the telson), lacking dorsal median carina (keels running down the center of the dorsal side), is laterally expanded. [2]
Other than the type species C. ventricosus, defined by its quadrate (square-shaped) pretelson and the narrow and elongated telson, one other species has been assigned to the genus; C. sarlei. The telson of C. sarlei is similar to that of C. ventricosus but the pretelson is shorter and wider. Both of these species have in the past been assigned to the larger and more derived pterygotid Pterygotus . [2] Some authors have speculated that if the chelicerae of Ciurcopterus are large (they are at present unknown from the genus), many fragmentary pterygotid specimens and species known only from chelicerae could be reassigned to Ciurcopterus. [2]
Ciurcopterus is classified as part of the pterygotid family of eurypterids, a group of highly derived eurypterids of the Silurian to Devonian periods that differ from other groups by a number of features, perhaps most prominently in the chelicerae (the first pair of limbs) and the telson. The chelicerae of the Pterygotidae were enlargened and robust, clearly adapted to be used for active prey capture and more similar to the claws of some modern crustaceans, with well developed teeth on the claws, than to the chelicerae of other eurypterid groups. [6] Another feature distinguishing the group from other eurypterid groups were their flattened and expanded telsons, likely used as rudders when swimming. [7] Their walking legs were small and slender, without spines, [8] and they were likely not capable of walking on land. [5]
Ciurcopterus was first described as a species of Pterygotus, P. ventricosus, by Erik. N. Kjellesvig-Waering in 1948. This species was represented by the dorsal impression of a single incomplete fossilised individual discovered near Kokomo, Indiana. The specimen (USNM 88130, currently housed at the U.S. National Museum in Washington) preserves most of the body, excluding parts of the appendages and the end of the telson. This individual would have measured approximately 29 cm (11 in) in length in life. Kjellesvig-Waering noted that the species did not resemble any of the other North American species of Pterygotus that had been described but that it did share some similarities with the British P. anglicus, from which it could still be differentiated by the different shape of the carapace, differences in the sixth appendage and P. ventricosus exhibiting a greater gibbosity. [9]
In 2007, O. Erik Tetlie and Derek E. G. Briggs redescribed the species based on four new specimens recovered from Kokomo. The new material allowed them to determine that P. ventricosus represented the most basal pterygotid eurypterid and the study helped provide evidence for the precise phylogenetic position of the family, showing that the Slimonidae (and not the Hughmilleriidae) was the most closely related group to the Pterygotidae. The specimens included YPM 208028 (the anterior half of an individual), YPM 209622 (a telson), YPM 210975 (a genital operculum) and YPM 210974 (a pretelson). [2] Tetlie and Briggs erected a new genus due to the unique features and distinct phylogenetic position of the species, naming it Ciurcopterus in honour of Samuel J. Ciurca, Jr., who has contributed significantly to eurypterid research by discovering a large amount of eurypterid specimens, including the four new specimens used to describe Ciurcopterus itself. [2] Another species, C. sarlei (also previously classified as a species of Pterygotus) was also referred to the genus due similarities in the pretelson (which is wider and shorter than that of C. ventricosus) and telson.
Ciurcopterus has been noted as possessing a mix of features from both more primitive pterygotioids such as Slimonia and more derived members of the group firmly within the Pterygotidae. For instance, its appendages share striking similarities with those of Slimonia but its carapace and more importantly its undivided genital appendage (a characteristic of pterygotid eurypterids) places it within the Pterygotidae. [2] C. ventricosus, also referred to as the "Kokomo pterygotid" after the site of its discovery, does possess some smaller differences compared to the rest of the Pterygotidae, such as keels running down the dorsal side of the pretelson. The enlarged chelicerae and claws present in all other pterygotids are unknown in Ciurcopterus since they have as of yet not been preserved in fossils, though this does not rule out that it had them. The combination of features in Ciurcopterus demonstrates that all the diagnostic characteristics of the Pterygotidae did not appear at the same time, suggesting that the evolution of these features was gradual. [2]
The cladogram below is based on the nine best-known pterygotid species and two outgroup taxa ( Slimonia acuminata and Hughmilleria socialis ). The cladogram also contains the maximum sizes reached by the species in question, which have been suggested to possibly have been an evolutionary trait of the group per Cope's rule ("phyletic gigantism"). [5] [10]
Pterygotioidea |
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
The Late Silurian Kokomo limestone that yielded the known fossils of C. ventricosus has also preserved fossil remains of numerous other eurypterid species and genera, including Erieopterus limuloides , Carcinosoma newlini , Onychopterella kokomoensis and Kokomopterus longicaudatus . [11] Fossils of various other organisms have also been recovered, including algal stromatolites, corals (such as Halysites ), small cephalopods (such as Protokionoceras ) and leperditiid ostracods. Two species of the conodont Spathognathodus (S. eosteinhornensis and S. snajdri) have also been recovered from the formation. [12]
The presence of stromatolites, molds of evaporate crystals and other features suggest that the Kokomo formation was primarily composed of very shallow environments. [12] Geological features of the formation, such as the argillaceous (resembling clay) limestone, suggests that the Silurian environment of the region might have been quiet and lagoonal. The bottom conditions were possibly anoxic and the environment might have been supratidal and hypersaline at large. [11]
Reconstructing the precise ecological role of Ciurcopterus may prove difficult as studies on the paleoecology of other pterygotid eurypterids have mainly focused not only on how visually acute they would have been in life, but also on the morphology of their claws and chelicerae, which are lacking in the known fossil Ciurcopterus specimens. Though derived pterygotids, such as Acutiramus, Jaekelopterus and Pterygotus, had divergent and specialized ecological roles, more basal genera, such as Erettopterus, were more generalized predators. [13]
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.
Pterygotus is a genus of giant predatory eurypterid, a group of extinct aquatic arthropods. Fossils of Pterygotus have been discovered in deposits ranging in age from Middle Silurian to Late Devonian, and have been referred to several different species. Fossils have been recovered from four continents; Australia, Europe, North America and South America, which indicates that Pterygotus might have had a nearly cosmopolitan (worldwide) distribution. The type species, P. anglicus, was described by Swiss naturalist Louis Agassiz in 1839, who gave it the name Pterygotus, meaning "winged one". Agassiz mistakenly believed the remains were of a giant fish; he would only realize the mistake five years later in 1844.
Slimonia is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Slimonia have been discovered in deposits of Silurian age in South America and Europe. Classified as part of the family Slimonidae alongside the related Salteropterus, the genus contains three valid species, S. acuminata from Lesmahagow, Scotland, S. boliviana from Cochabamba, Bolivia and S. dubia from the Pentland Hills of Scotland and one dubious species, S. stylops, from Herefordshire, England. The generic name is derived from and honors Robert Slimon, a fossil collector and surgeon from Lesmahagow.
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.
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.
Acutiramus is a genus of giant predatory eurypterid, an extinct group of aquatic arthropods. Fossils of Acutiramus have been discovered in deposits of Late Silurian to Early Devonian age. Eight species have been described, five from North America and two from the Czech Republic. The generic name derives from Latin acuto and Latin ramus ("branch"), referring to the acute angle of the final tooth of the claws relative to the rest of the claw.
Jaekelopterus is a genus of predatory eurypterid, a group of extinct aquatic arthropods. Fossils of Jaekelopterus have been discovered in deposits of Early Devonian age, from the Pragian and Emsian stages. There are two known species: the type species J. rhenaniae from brackish to fresh water strata in the Rhineland, and J. howelli from estuarine strata in Wyoming. The generic name combines the name of German paleontologist Otto Jaekel, who described the type species, and the Greek word πτερόν (pteron) meaning "wing".
Nanahughmilleria is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Nanahughmilleria have been discovered in deposits of Devonian and Silurian age in the United States, Norway, Russia, England and Scotland, and have been referred to several different species.
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.
Salteropterus is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Salteropterus have been discovered in deposits of Late Silurian age in Britain. Classified as part of the family Slimonidae, the genus contains one known valid species, S. abbreviatus, which is known from fossils discovered in Herefordshire, England, and a dubious species, S. longilabium, with fossils discovered in Leintwardine, also in Herefordshire. The generic name honours John William Salter, who originally described S. abbreviatus as a species of Eurypterus in 1859.
Erettopterus is a genus of large predatory eurypterid, an extinct group of aquatic arthropods. Fossils of Erettopterus have been discovered in deposits ranging from Early Silurian to the Early Devonian, and have been referred to several different species. Fossils have been recovered from two continents; Europe and North America. The genus name is composed by the Ancient Greek words ἐρέττω (eréttō), which means "rower", and πτερόν (pterón), which means "wing", and therefore, "rower wing".
Pterygotidae is a family of eurypterids, an extinct group of aquatic arthropods. They were members of the superfamily Pterygotioidea. Pterygotids were the largest known arthropods to have ever lived with some members of the family, such as Jaekelopterus and Acutiramus, exceeding 2 metres (6.6 ft) in length. Their fossilized remains have been recovered in deposits ranging in age from 428 to 372 million years old.
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.
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 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.
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.
Slimonidae is a family of eurypterids, an extinct group of aquatic arthropods. Slimonids were members of the superfamily Pterygotioidea and the family most closely related to the derived pterygotid eurypterids, which are famous for their cheliceral claws and great size. Many characteristics of the Slimonidae, such as their flattened and expanded telsons, support a close relationship between the two groups.
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.
Necrogammarus salweyi is the binomial name applied to an arthropod fossil discovered in Herefordshire, England. The fossil represents a fragmentary section of the underside and an appendage of a pterygotid eurypterid, a group of large and predatory aquatic arthropods that lived from the late Silurian to the late Devonian. The Necrogammarus fossil is Late Silurian in age and its generic name means "dead lobster", deriving from Ancient Greek νεκρός and Latin gammarus ("lobster").
This timeline of eurypterid research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of eurypterids, a group of extinct aquatic arthropods closely related to modern arachnids and horseshoe crabs that lived during the Paleozoic Era.