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Onychopterella

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.

Description
Like the other onychopterellids, Onychopterella was a small eurypterid. The smallest species, O. pumilus, measured only . The size of the largest one, O. kokomoensis, is estimated at , representing the biggest species of the family Onychopterellidae. O. augusti had a similar size, with the largest specimen reaching . The prosoma (head) had a subquadrate (almost square) shape in O. kokomoensis The outline of the carapace (the exoskeleton part covering the prosoma) varied according to the species. It was anteriorly rounded in the corners in O. kokomoensis, and O. pumilus, the eyes were reniform (kidney-shaped), standing out for their size and prominence in the latter species. ==History of research==
History of research
In 1896, paleontologists Samuel Almond Miller and William Frank Eugene Gurley described a new species of Eurypterus, E. kokomoensis, based on four specimens, three of them well-preserved and a fragmentary one, FMNH UC6638, an almost complete specimen, was designated as the type specimen. They also described E. ranilarva as a close relative of O. (Eurypterus) kokomoensis, differing from the latter mainly in the greater width of the carapace. In 1951, paleontologist and geologist Leif Størmer noticed that the name Onychopterus had been used previously by a bird genus introduced in 1850 by the ornithologist Ludwig Reichenbach. Størmer substituted the old name with a new one, Onychopterella. In 1916, researcher and geologist Thomas Edmund Savage erected the new species Eurypterus pumilus to accommodate one single well-preserved specimen showcasing the ventral side of the body It has been debated whether O. pumilus really belongs to this genus. In 1999, professor and paleobiologist Roy E. Plotnick considered that the species represented a form of Drepanopterus, while geologist and paleobiologist James C. Lamsdell suggested in 2012 that it belonged to Stoermeropterus because of the similarities of the metastoma, genital appendage and telson. A 1995 study led by paleontologist Simon J. Braddy proposed that because of its small size, O. pumilus could simply represent an ontogenetic stage (a different developmental stage of the same animal throughout its life) of O. kokomoensis. Due to the lack of a known accession number for the fossil, its re-examination is impossible. In 1995, paleontologists Braddy, Richard John Aldridge and Johannes N. Theron described a well-preserved eurypterid from the Soom Shale Member of the Table Mountain Sandstone, Cape Province, South Africa, and named it O. augusti. It is based on two specimens of which one preserves soft tissues. The holotype (GSSA C373, housed at the Geological Survey of South Africa in Pretoria along with the paratype) was discovered by August Patrick Pedro, honoured in the specific epithet augusti. It differed from the rest of the species by the lack of large epimera in the pretelson, wider body proportions, the short length of the postabdomen and telson, the lanceolate form of the latter, the two projections of the eighth podomere and in a distal spine longer than in the rest of the species. The paratype, GSSA C427, is the largest known specimen. O. augusti was also compared to the enigmatic Silurian eurypterid Marsupipterus sculpturatus, concluding that the differences between the telson (the only known part of Marsupipterus) of both species are probably preservational. ==Classification==
Classification
E. ranilarva), housed at the Field Museum of Natural History, Chicago Onychopterella is classified as part of its own family, Onychopterellidae, the only clade (taxonomic "group") within the monotypic superfamily Onychopterelloidea. It was originally described as a subgenus of Eurypterus, In 2011, Lamsdell recovered Onychopterella as monophyletic (a group consisting of all the descendants of the last common ancestor, i.e. a valid genus), considering Moselopteroidea the most basal eurypterine clade. He also erected the new superfamily Onychopterelloidea and family Onychopterellidae, placing within the latter the genera Onychopterella and Tylopterella. This family was characterized by the presence of spines in the second to fourth pairs of appendages, lack of spines in the fifth and sixth (except occasionally a distal spine in the last podomere of the sixth appendage), the shape of the carapace with lateral eyes and a lanceolate or styliform telson, among other features. Alkenopterus was assigned to Onychopterellidae three years later because of the detection of a movable spine in the swimming leg, rather than a simple projection as previously thought. Before the creation of Onychopterellidae, Onychopterella had been classified in the family Erieopteridae since 1989 by paleontologist Victor P. Tollerton, initially together with Erieopterus and Buffalopterus based on similarities of the morphology of the appendages and the opisthosoma. The cladogram below is based on a larger study (simplified to only show eurypterids) in a 2011 phylogenetic analysis carried out by Lamsdell, showcasing the basal members of the Eurypterina suborder of eurypterids with other derived groups. '', a closely related onychopterellid genus }} ==Paleobiology==
Paleobiology
view of the best preserved book gill of O. augusti'' Only the alimentary canals of a few species of eurypterids, such as Carcinosoma newlini, Acutiramus cummingsi and Eurypterus lacustris, have been described. However, the excellent levels of preservation of the O. augusti fossils have allowed the description of what most likely was the anterior portion of the gut. Between the two coxal (from the coxa, the point of union with the appendages) muscle masses of the swimming leg, a spiral-shaped structure was found. A similar structure in the stylonurine Hibbertopterus wittebergensis, in which a spiral valve has been found near the posterior zone of the prosoma, has been reported. These valves have developed independently in certain groups of fishes in response to the need to increase the absorptive area of the gut, so Braddy, Aldridge and Theron considered the case was the same for the eurypterids. Nevertheless, the presence of this valve may have depended on the digestion and absorption requirements of each eurypterid, in turn depending on factors such as the feeding habitats or the size of the animal. The appendages of H. wittebergensis suggest it was a sediment feeder, using its valve to extract food from the soil and increase the absorptive capacity of the gut. It is possible that the spiral structure of O. augusti had a similar function, acting as a valve for the mesenteron (midgut, formed by the stomach and intestines), thus increasing the absorptive area of this region. Apart from this spiral structure, the pretelson of the paratype preserves a faint impression that expanded posteriorly to a depression in the medial area, representing an anal opening. Its great preservation has also allowed the examination of the internal muscular structure of the swimming leg. The coxal muscles were elongated towards an anterolateral orientation. This suggests the legs were maintained in a posterolateral position to walk in order, thus allowing sufficient stability at the animal's center of gravity, The eurypterid ichnospecies (a species based on fossil prints) Palmichnium capensis is thought to have belonged to O. augusti. This ichnospecies was found in the Table Mountain Sandstone and dates from the Ordovician, coinciding with O. augusti. Its trackways were medium-sized (largest track wide) and consisted of several symmetrical series of four tracks and individual typically oval or tear-shaped marks with small impressions on the sides, sometimes bilobed and intermittent. A median line was occasionally displayed. Several factors support that P. capensis represents the tracks of O. augusti; the average external width of P. capensis is comparable to that of fossils of O. augusti, the distal spines of the swimming leg of O. augusti could have caused such bilobed marks and Onychopterella represents the only Ordovician eurypterid from those deposits. In addition, the median line could have been produced by the telson touching the substrate. O. augusti would have been able to make incursions to the terrestrial surface, but it would have been uncomfortable for it, performing an undulatory gait and keeping its telson in regular contact with the ground, hence the median line. But nevertheless, this can not be completely proven unless a specimen of O. augusti is found alongside similar tracks. ==Paleoecology==
Paleoecology
It is thought that the genus Onychopterella lived in brackish (water with salt but not completely salty) and possibly marine water. It was undoubtedly nektobenthic, that is, being able to swim but staying most of the time in the stratum. The type species, O. kokomoensis, inhabited a peritidal environment (in a part of the shore) along with other eurypterids such as Drepanopterus longicaudatus, Carcinosoma newlini and Erieopterus limuloides, as well as with the ostracod Leperditia ohioensis and the brachiopods Pentamerus divergens and Schuchertella interstriata. It was Early Pridoli (Late Silurian) in age, living in Indiana. On the other hand, O. pumilus, which lived in the Early Llandovery (Early Silurian) epoch of Illinois, is associated with many indeterminate species of anthozoans, craniates, brachiopods and gastropods in an open shallow subtidal (the sunlight reaches the bottom of the ocean) environment. Most of the eurypterids found outside the supercontinent Laurentia were highly derived genera that lived during the prelude of the amalgamation (union) of Pangea, as well as genera with a more effective morphology for dispersion. Therefore, the isolated appearance of an ancient form like O. augusti in the supercontinent Gondwana is challenging for the scientific community to explain, since the genus was not a good disperser due to the narrow form of its swimming legs compared to other more derived groups such as Pterygotidae or Adelophthalmidae. It seems unlikely that the eurypterines originated in Gondwana since there is no record of fossil remains of stylonurines or other basal swimming forms. It also seems that the collectivity of O. augusti went extinct without leaving a viable population of eurypterids in Gondwana. It has been proposed that O. augusti established itself in Gondwana by crossing the sea floor during the periods of sea level descent of the Hirnantian. Onychopterella does not represent the only occurrence of basal genera in Gondwana; Paraeurypterus, a genus known from deposits of the Şort Tepe Formation in southeastern Turkey, probably arrived there by the same method as O. augusti. ==See also==
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