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Megistaspis

Megistaspis is a genus of asaphid trilobites that lived throughout the Early and Middle Ordovician. Megistaspis was common throughout the Early and Middle Ordovician of Baltoscandia, but specimens have also been found in Australia, France, Germany, Morocco, and the United States. Appendages and a digestive system of M. hammondi have been preserved; it likely served as a mixed detritivore, producing Cruziana rugosa trace fossils. Meanwhile, Megistaspis hyorrhina may have burrowed below the sediment, using a swelling on the head to detect pressure changes. Megistaspis is also notable as it displays a large degree of morphological variation as a result of environmental conditions like ocean depth and substrate. The genus plays an important part in the biostratigraphy of Ordovician Baltoscandia, with several biozones being named after Megistaspis species.

History
The type species of Megistaspis, M. (Megistaspis) limbata, was described as Trilobites limbatus by Norwegian zoologist Christian Boeck in 1838. In 1851, Swedish paleontologist Nils Peter Angelin moved Trilobites limbatus, along with Entomostracites extenuatus and Asaphus heros, to the new genus Megalaspis and erected several new species, including Megalaspis gigas, Megalaspis explanata, and Megalaspis planilimbata. In 1956, Estonian-Swedish geologist Valdar Jaanusson recognized that the name Megalaspis was preoccupied by the fish genus Megalaspis, named by Pieter Bleeker earlier in 1851. He therefore erected the genus Megistaspis to replace Megalaspis, choosing the name, which is Greek for "largest shield", to be as similar to the old name Megalaspis as possible. Jaanusson created two subgenera: M. (Megistaspis) to accommodate forms with a strongly convex thoracic rachis and triangular cephalon, and M. (Megistaspidella) to accommodate later forms with a flat thoracic rachis and elongate cephalon, as well as suggesting the existence of a third group, composed of early forms with a somewhat flat thoracic rachis but semicircular cephalon, which he called the M. planilimbata group. Also in 1956, Scandinavian paleontologist Torsten E. Tjernvik moved Megistaspis planilimbata to the subgenus Plesiomegalaspis (Plesiomegalaspis) and erected the new species Plesiomegalaspis estonica, Plesiomegalaspis norvegica, and Plesiomegalaspis scutata in addition to creating the new subgenus Plesiomegalaspis (Ekeraspis), containing forms with a long pygidial spine and deep posterior border furrow of the cephalon, to accommodate Megalaspis heroides and his new species Plesiomegalaspis armata. In 1976, Soviet paleontologist E. A. Balashova split the genus Megistaspis into several genera as part of the family Megistaspisidae, which also included Plesiomegalaspis and several other genera. Balashova (1976) created several new genera: she created the genus Paramegistaspis, comprising forms previously within the M. planilimbata group, to accommodate P. planilimbata, P. estonica, P. norvegica, P. scutata and their relatives and the genus Rhinoferus, composed of forms with a large swelling on the glabella, to accommodate M. hyorrhina and its relatives. In addition, M. (Megistaspidella) was split into the genus Megistaspidella and P. (Ekeraspis) was split into the genus Ekeraspis. In 1983, B. T. Wandås created the subgenus M. (Heraspis), containing forms with a wide cephalon and a long pygidial spine, to accommodate two species: M. heroica and M. laticauda. In 1995, Dutch paleontologist Arne Thorshøj Nielsen demoted the genera Megistaspidella, Paramegistaspis, Rhinoferus, and Ekeraspis to subgenera of Megistaspis, giving Megistaspis six subgenera: Megistaspis, Megistaspidella, Paramegistaspis, Rhinoferus, Ekeraspis, and Heraspis. Heraspis was subsequently subsumed into Megistaspidella by Hansen, 2009. == Description ==
Description
Megistaspis is a large genus of asaphid trilobite, with several species reaching lengths in excess of 25 centimeters. The pygidium is roughly equal in size to the cephalon, is triangular or semicircular, and, like the cephalon, has concave lateral parts that form a distinct marginal rim. Appendages Preserved appendages have been found in several specimens of M. hammondi, a large species of Megistaspis from the Fezouata Biota. One specimen in particular, MGM-6756X, preserves a complete set of endopods. This specimen shows that M. hammondi has 21 pairs of limbs: three pairs of cephalic limbs, eight pairs of thoracic limbs, and 10 pairs of pygidial limbs. M. hammondi has slight heteropody as its cephalic limbs are larger and heavier than its thoracic and pygidial limbs. The cephalic limbs also bear spines on podomeres 2-4, with the best-preserved cephalic appendage of specimen MGM-6756X preserving 11 spines dorsally and 7 spines ventrally. No spines are present on the thoracic or pygidial limbs. An additional specimen, MGM-7569X, preserves part of the right antenna in addition to the distal portions of the endopods and exopods. Antennae can also be observed in many commercially sold specimens, but they usually exhibit varying degrees of cosmetic modification, including the addition of extra podomeres, the adding of spines to the abaxial side of the antenna, and the outlining of the rock surrounding the antennae with "cat ears". s of M. hammondi specimen MGM-6756X|center Digestive system s Another specimen of M. hammondi, MGM-6755X, preserves the digestive system. The alimentary canal of MGM-6755X consists of an 8-millimeter wide crop that extends from the anterior edge of the eyes to the end of the cephalon, tapering to 4 millimeters by the end. Following the crop is a 3-millimeter wide intestine that is preserved up to the third thoracic segment before reappearing for 22 millimeters before the axial end of the pygidium. There are two bilaterally symmetrical digestive caecae anterior to the crop that occupy the whole anterior region below the glabella. Behind the eyes are at least 4 additional pairs of lobed, laterally-oriented caecae with longitudinally aligned tips. The posterior caecae lengthen transversely as the crop tapers. The first three thoracic segments preserve digestive caecae similar in size and shape to the posteriormost cephalic pair. The alimentary canal may preserve three pairs of small and simple caecae posterior of the pygidial axis; however, the presence of caecae along the entire digestive tract is impossible to determine due to the quality of preservation and the fact that much of the specimen is restored. The crop and intestine are preserved with a positive relief, while the caecae are preserved as voids but were likely initially permineralized during early diagenesis, possibly due to being an enzymatically active region in life. The presence of both a crop (anatomy) and caecae suggests that M. (Ekeraspis) hammondi had a unique "type 3" digestive system (in contrast to the "type 1" digestive system defined by the presence of caecae but no crop and the "type 2" digestive system found in Isotelus and Birmanites defined by the presence of a crop but no caecae). Ornamentation The test of Megistaspis is generally smooth. Terracing exists only on the doublure (only on the base of the "snout" in forms like M. acuticauda with elongated snouts), the articular facets of the thorax and pygidium, and the genal spines, while light terracing may exist along the margins of the free cheeks and pygidium. However, small pits, which are indicated on the internal surface of the test by small tubercles, occur in all Megistaspis species (but are only preserved in cases where the test has not been weathered or corroded). The pits may be in close proximity to each other or widely spaced from each other depending on the species; in addition, the pits are not always evenly spread across the test and may be crowded around the borders of the test but widely spaced closer to the central regions. In some cases, the pits may be of two different sizes, and in these cases, the smaller pits form a background. Below the surface of the test is a perfectly smooth lamella which can be observed in specimens where the surface of the test is peeled off. The pits are clearly visible on the lamella and can also be seen on the mold if the sediment is fine enough to preserve almost microscopic details. In some preserved pygidia with a weathered test, fine striations can be seen diverging forwards and outwards to the dorsal furrow, where they suddenly turn outwards and somewhat backwards. This structure is entirely independent of the relief of the pygidium, and the weak ribs in the posterior parts of the pygidium can be superseded by the striations. In the anterior ribs, the distal portion of the ribs behind the rib furrow may become diffuse and merge into the network created by the striae. The striations are preserved in both external and internal relief and can sometimes be preserved in an internal mold of the pygidium. The striations are also not restricted to the pygidium, being observed in cranidia of M. (Rhinoferus) lawrowi and M. (Megistaspidella) heros. These striations have been found in M. (Megistaspidella) heros, M. (Megistaspidella) acuticauda, M. (Megistaspidella) curvispina, and M. (Megistaspidella) gigas, but indications of them can be seen in other species and they likely exist in all species of Megistaspis and perhaps other asaphid genera. == Paleobiology ==
Paleobiology
Diet and trace production The relief of the crop and intestine of M. (Ekeraspis) hammondi has the same texture and grain size as the matrix, suggesting active sediment ingestion or rapid sediment infill. This, combined with the slight heteropody of the cephalic appendages (where the cephalic limbs were slightly larger than the thoracic limbs) and the non-forked hypostome, suggests that M. (Ekeraspis) hammondi was a detritus feeder, a common mode of life for other benthic trilobites. The spinose cephalic appendages of M. (Ekeraspis) hammondi were likely used to dig for food in a comb-like motion. This behavior would have produced Cruziana rugosa traces, which were pascichnia (combined feeding and locomotion traces) associated with the food searching strategy of trilobites. In addition, Cruziana rugosa traces could have reached 26 centimeters wide, which is only consistent with large asaphoid trilobites like M. (Ekeraspis) hammondi and Ogyginus, the latter of which are frequently found in association with Cruziana rugosa traces. In addition, a specimen of M. (Paramegistaspis) planilimbata cyclopyge from Montana shows a scarred glabella, likely due to parasitism. Boring organisms Megistaspis exuviae from the Vaginatum limestone are rapidly destroyed by seawater and boring organisms if they are not buried immediately after molting. The surfaces of eroded tests frequently bear branching, winding, and anastomosing ridges with a semicircular cross-section without any structures like pores or striations. These ridges are never seen in specimens with smooth tests and likely contributed to the destruction of the shell. While the organism that produced these tubes may never be discovered, a colony of bryozoans has also been seen on a heavily eroded specimen, proving that the bioerosion was caused by an organism contemporaneous with Megistaspis. == Species ==
Species
Megistaspis ringsakerensis Skjeseth, 1952 • Megistaspis knyrkoi Schmidt, 1906''' This subgenus is characterized by a triangular cephalon and pygidium, a notched hypostome, a glabella with lateral grooves, basal lobes and occipital lobes, a convex rachis and pleurae, the presence of a postmarginal furrow, and a pygidial rim that widens slightly towards the posterior end of the pygidium. • Megistaspis (Megistaspidella) grandis Sars, 1835 – NorwayMegistaspis (Megistaspidella) isvosica Balashova, 1976 • Megistaspis (?Megistaspidella) laine Jaanusson, 1956 – Volkhov stage of SwedenMegistaspis (Megistaspidella) lamanskii Schmidt, 1906 • Megistaspis (Megistaspidella) heros Dalman, 1828 • Megistaspis (Paramegistaspis) popovkiensis Balashova, 1966 – Early Ordovician of RussiaMegistaspis (Paramegistaspis) putilovensis Balashova, 1966 – Arenig of RussiaMegistaspis (Paramegistaspis) scutata Tjernvik, 1956 – Arenig of Sweden • Megistaspis (Rhinoferus) ?hyorrhina var. kolenkoi Schmidt, 1906 – Norway This subgenus is characterized by a cephalon with long genal spines, a subtriangular pygidium with a long terminal spine, a short hypostome with a rounded posterior margin, a glabella without lateral grooves and occipital lobes, a convex rachis and pleurae, a wide and deep postmarginal furrow, and a narrow rhachis. • Megistaspis (Ekeraspis) hammondi Brögger, 1882 – Tremadocian of MoroccoMegistaspis (Ekeraspis) heroides Brögger, 1882 – Arenig of Norway and Sweden == Biostratigraphy ==
Biostratigraphy
The genus Megistaspis is one of the four most common genera in the of Sweden, (the others being Nileus, Symphysurus, and Geragnostus), and its abundance and diversity allows it to be used as an index fossil for several biozones (rock layers defined by the presence of a certain species of index fossil) of Scandinavia and Estonia. • the M. (Ekeraspis) armata, M. (Paramegistaspis) planilimbata, and '''M. aff. (Paramegistaspis) estonica''' zones of the Ottenbyan stage. • the M. (Paramegistaspis) estonica zone of the Billingenian stage. • the M. (Megistaspis) polyphemus, M. (Megistaspis) simon, and M. (Megistaspis) limbata zones of the Volkhovian stage. • the M. (Megistaspidella) gigas-M. (Megistaspidella) obtusicauda zone of the stage. == References ==
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