Megalograptus
Megalograptus is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of "Megalograptus" have been recovered in deposits of Katian (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 "great writing" and originates from the mistaken original belief that "Megalograptus" was a type of graptolite, often given names ending with "-"graptus"" (writing). "Megalograptus" was a large predatory  megalograptid eurypterid, with the largest and best known species, "M. ohioensis", reaching lengths of . Some species were substantially smaller, with the smallest, belonging to a hitherto undetermined species, only growing to about in length. Morphologically, "Megalograptus" was highly distinct. The two most distinctive features of "Megalograptus" were its massive and spined forward-facing appendages, far larger than similar structures in other eurypterids, and its telson (the last division of the body). The sharp spike-shaped telson of "Megalograptus" was not venomous, but it was specialized in that it was surrounded by unique cercal blades, capable of grasping. Certain fossils of  three different species, "M. ohioensis", "M. shideleri" and "M. williamsae", are so well-preserved that researchers have been able to infer the coloration they might have possessed in life. All three were deduced to have been brown and black in color, with "M. ohioensis" being darker than the others. First described by Samuel Almond Miller in 1874, based on fragmentary fossil remains of the species "M. welchi", "Megalograptus" being a graptolite was not formally questioned until 1908, when Rudolf Ruedemann recognized the fossils as eurypterid remains. "Megalograptus" was noted as being similar to "Echinognathus" by August Foerste in 1912 and  the two genera have been considered closely related since then, and have been grouped together in the Megalograptidae since 1955. In 2015, the genus "Pentecopterus" was also assigned to the family. Kenneth E. Caster and Erik N. Kjellesvig-Waering revised "Megalograptus" in 1955, owing to the discovery of more complete fossil material of the new species "M. ohioensis". Caster and Kjellesvig-Waering conducted further work on "Megalograptus" over the following years. In 1964, they named the species "M. shideleri" and "M. williamsae" and reclassified the fragmentary eurypterid "Ctenopterus alveolatus" as a species of "Megalograptus". "Megalograptus" lived in near-shore marine  environments, where it used its large appendages, and possibly its telson and cercal blades, to capture prey. Possible coprolites (fossilized dung) are known from "M. ohioensis", which contain fossil trilobite fragments as well as fragments of "M. ohioensis" itself. This suggests that "Megalograptus" might have been cannibalistic at times, like many modern chelicerates. Description. Size. "Megalograptus" was a genus of large megalograptid eurypterids. The largest species was "M. ohioensis", which ranged in length from . "M. ohioensis" was the second largest megalograptid and the second largest eurypterid of the Ordovician period, smaller only than the related "Pentecopterus", which could grow to  . Previous estimates have placed the size of some species of "Megalograptus" as substantially larger, with the type species "M. welchi" once believed to have reached lengths of in length. According to a 2009 study by James Lamsdell and Simon J. Braddy, such estimates are dubious as they are based on ornamentation in incomplete fossils. In the case of "M. shideleri", once estimated to have reached lengths of , the size estimate was based only on two fragmentary tergites (upper portions of body segments), wherein the dimensions of the ornamental scales were unusually large, interpreted as suggesting a giant body size. The  fact that scales can vary in size across the bodies of megalograptid eurypterids and that one of the relevant tergites of "M. shideleri" was not longer than 3 cm (1.2 in) suggests that this species did not reach lengths of more than . The length of the species "M. alveolatus" is uncertain, but it was much smaller than "M. ohioensis", and "M. williamsae" grew to about . The smallest known species of "Megalograptus" was an as yet undescribed Canadian species which only grew to 10 cm (3.9 in) in length. Morphology. "Megalograptus" was morphologically unique and easily distinguishable from other eurypterids. The  carapace (head plate) of "Megalograptus" was vaguely quadratic in shape and flattened, lacking a marginal rim, which was present in some other eurypterids. At the front of the carapace there was a downturn and six small downward-facing spikes, possibly an adaptation for digging in the mud. The compound eyes of "Megalograptus" were medium-sized and reniform (kidney-shaped), located close to the edge of the carapace. The ocelli (simple eyes located more centrally) were small. Some of the appendages of "Megalograptus" unusually had one more joint than was common in eurypterids. Among the appendages, the third pair (counting the  simple chelicerae, pincers or mouth parts, as the first pair) are the most notable. In "Megalograptus" these were massive structures, covered in pairs of great spines, only comparable to the same structures in "Mixopterus", another eurypterid. The appendages of "Megalograptus", about 3.5 times the length of the carapace, were significantly larger than those of "Mixopterus". On the fourth joint of the appendages, one pair of spines end in bulbous structures, rather than sharp points, and were perhaps sensory. The fourth pair of appendages were short and spiny, but the fifth pair, immediately preceding the swimming paddles (placed on the sixth  and final pair of appendages), were completely spineless. This unusual limb is similar to the same appendage in the distantly related genus "Eurypterus", where it has been interpreted as a balancing organ. The third joint of the swimming paddles of "Megalograptus" bent the appendages forwards, a rare feature in the eurypterids, otherwise mostly known from the distantly related genus "Dolichopterus". The wide swimming paddles of "Megalograptus" were formed from the sixth joint of the appendage. The seventh joint, which in many genera formed a major part of the paddle, was reduced to a relatively small structure. The eighth joint, not  preserved in any known "Megalograptus" fossil material, is indicated as existing by attachment points in the seventh joint, which also indicate that it was significantly smaller than in other eurypterids.The mesosoma of "Megalograptus" (the first six segments after the head) was distinctly similar to the same segments in modern scorpions and different from the mesosomas of other eurypterids. The body contracted after the last segment of the mesosoma, rather than after the first segment of the metasoma (the last six segments), which was otherwise typical for eurypterids. In most eurypterids, the mesosoma was widest at the fourth or fifth  segment, but in "Megalograptus" it was widest at the third. The first segment of the mesosoma was considerably shorter than the succeeding segments, which were otherwise approximately of the same length. The last few segments of the body were slightly longer than the preceding segments. The metastoma (a large plate located on the underside of the body) of "Megalograptus" was roughly egg-shaped, unusually wide and broadly subtriangular (almost triangular) in shape, differentiating it from the same structure in all other eurypterids, where it was usually cordate (heart-shaped). The most unusual feature of "Megalograptus" was the structure formed by  the telson (the last division of the body) and the immediately preceding segment (the pretelson). "Megalograptus" had, alongside the sharp and stout telson spike, two paired and rounded blade-formed lobes, the so-called cercal blades. These were attached beneath the telson, directly to the pretelson. The blades were capable of articulation, in effect forming a large grasping organ. In other eurypterids, the telson tends to be an undivided structure in the shape of a paddle or spike, meaning that the cercal blades distinguish "Megalograptus" from nearly all other eurypterids. Cercal blades are only known from one other eurypterid, "Holmipterus , and are lacking in the basal (primitive") megalograptid "Pentecopterus". Taken together with the telson, the telson–pretelson assemblage of "Megalograptus" forms a flattened structure, superficially similar to the flattened telsons of many genera in the superfamily Pterygotioidea. "Megalograptus" was ornamented with small scales of irregular size across its body. On the carapace, they were flat and disc-like and scattered without any obvious pattern. On the fifth pair of appendages, the scales were more elongated. On the main body, the scales were rounded, raised and nearly elliptical in shape. Many of the scales on the carapace, the fifth pair of  appendages, the mesosoma and metasoma and some on the appendages had holes in their center, suggesting that they once supported bristles (stiff hairs). In life, "Megalograptus" may have had an almost hirsute (hairy) appearance. Coloration. In the 1964 description of "M. ohioensis", "M. shideleri" and "M. williamsae", Kenneth E. Caster and Erik N. Kjellesvig-Waering made inferences of the life coloration of the species based on a collection of well-preserved specimens. In some fossils of "M. shideleri", the fossils retained their original coloration, with no replacement having taken place (meaning that mineralization during fossilization did not distort the original  color scheme of the fossils). "M. shideleri" was brown, with scales varying in color from dark brown to black and the integument (the scales) was a lighter brown color. A similar, but darker, brown and black color scheme has been inferred for "M. ohioensis", and its fossils being better preserved allows for more detailed examination. "M. williamsae" also had a similar color scheme, with its tergites indicating black scales against light brown integument. The colors of "Megalograptus" inferred by Caster and Kjellesvig-Waering are similar to those inferred by Kjellesvig-Waering of "Carcinosoma newlini", another eurypterid also inferred to have  been brown and black, in 1958. According to Caster and Kjellesvig-Waering, "M. ohioensis" was mostly dark brown, with some black elements. Though no obvious integument pattern has been determined, most of the body, including the head, had dark brown integument, contrasted by black scales. The coxae (base segments of the appendages) were dark brown, with black scales and black gnathobases ("tooth plates" surrounding the mouth). As such, the appendages began as dark brown in color, but quickly darkened towards their ends. Most of the appendages of larger specimens, including the spiny and large forelimbs, were almost entirely black in  color and with black spines, although in smaller specimens, the appendages were typically lighter in color. Connective tissue in the appendages was pale brown in color. The appendage immediately preceding the swimming paddles was not entirely black, instead just darkening to a very dark brown. The metastoma was dark brown, with black scales, postules (small elevations) and mucrones (tiny spines). The telson of "M. ohioensis", as well as much of the preceding segment, was entirely black in color. History of research. Type material. Fossils of "Megalograptus" were first described by Samuel Almond Miller in 1874. Miller mistakenly believed the fossil  material, consisting of a postabdominal (segments 8–12) tergite and two fragments of an appendage, was the integument of a graptolite (a member of Graptolithina, an extinct group of colonial pterobranchs), and gave it the name "Megalograptus", meaning "great writing" (deriving from the Greek "megale", "great", and "graptos", "writing", commonly used for graptolite fossils). One reason for Miller's mistaken identification is that the exact outline of the fossils was unclear because they were not properly cleaned yet. The fragmentary fossils of "M. welchi" were initially recovered by L. B. Welch, whom the species name "welchi" honours, near Liberty, Ohio, in  rocks of Katian (Late Ordovician) age of the Elkhorn Formation. With the exception of the type material, "M. welchi" is only fragmentarily known. It is probable that more fossils could have been uncovered if it had been immediately recognized as a large eurypterid. By the time it was recognized as such and the fossils were deemed to be of interest, further work at the fossil site had irreversibly damaged what remained of the eurypterid fossils. The status of "Megalograptus" as a graptolite was first questioned in 1908 by Rudolf Ruedemann, who was researching Ordovician graptolites. Ruedemann instead recognized the remains  of "M. welchi" as eurypterid fossils. That same year, Ruedemann's suspicions were confirmed in discussions with August Foerste and Edward Oscar Ulrich, who also agreed that the fossils were of a eurypterid. Foerste was invited to contribute with his understanding of "Megalograptus" to Ruedemann's and John Mason Clarke's 1912 monograph "The Eurypterida of New York". He recognized "Megalograptus" as similar to the eurypterid "Echinognathus clevelandi", assuming them to either be related or of the same genus. The features uniting the two were noted to be the spines of the appendages, the scaly ornamentation and the longitudinal ridges  of the preserved segment. Foerste also noted that the fossils of "M. welchi" were not morphologically distinct enough from other eurypterids to differentiate it, with its earlier age instead serving as the main distinction of the genus and species. Additional fossils. "Megalograptus" was considerably revised in 1955 by Caster and Kjellesvig-Waering in Leif Størmer's 1955 "Treatise on Invertebrate Paleontology", from which the modern understanding of the genus originates. The revision was made possible with the discovery of new fossil material, consisting of what at the time was the best preserved Ordovician eurypterid fossil material discovered. These fossils, found  in deposits of Katian age alongside the Ohio River road (U.S. Route 52), approximately 14.5 kilometres (9 miles) north of Manchester, Ohio, were assigned to a new species, "M. ohioensis". The type material of "M. welchi" was compared to the new fossils by Caster and Kjellesvig-Waering, though only in the "walking legs" (i.e. the second to fifth pair of appendages) given that they were the only body part preserved for both "M. welchi" and "M. ohioensis". While recognized as being of the same genus, Caster and Kjellesvig-Waering also noted differences, supporting the species distinction of "M. welchi" and  M. ohioensis, including the leg of "M. welchi" being stouter, with thicker and shorter spines, and some differences in the joints (in "M. welchi", the second joint had spines, which it did not in "M. ohioensis", and in "M. welchi", the spine-shaped ultimate joint was blunt and thick, whereas it was slender in "M. ohioensis"). In 1964, Caster and Kjellesvig-Waering named two new species of "Megalograptus" based on additional fossil material from the Katian of Ohio: "M. shideleri" and "M. williamsae". "M. shideleri" was named based on fragmentary fossil specimens originally found by William H. Shideler in the  Saluda Formation near Oxford, Ohio, and in Indiana. The species is named in his honor. "M. shideleri" differs from "M. ohioensis" in that its gnathobases have fewer denticles and a much more developed second tooth. The "M. shideleri" fossils could not be compared to the type material of "M. welchi" as there is no overlap in the preserved body parts. "M. williamsae" was named based on a cercal blade, alongside fragments of tergites and appendages, discovered in the Waynesville Formation, near Clarksville, Ohio, by Carrie Williams, whom the species name honours. "M. williamsae" differs from "M. ohioensis" in its cercal  blades, with a slightly different pattern of scales and longer, narrower and sharper end points, rather than the more acute, hooked and stouter end points in "M. ohioensis". The species "M. alveolatus" was originally named as a species of the very distantly related "Ctenopterus" by Ellis W. Shuler in 1915, based on fossil fragments, including of the appendages and the telson spike, collected in Late Ordovician deposits along Walker Mountain in Virginia belonging to the Bays Formation. It was the first eurypterid to be described from Virginia. The species name "alveolatus" refers to the pronounced development of the alveolar processes  (pits) around the spines. Because of the fragmentary state of its fossils, "M. alveolatus" has had a complex taxonomic history. Although Kjellesvig-Waering initially believed that it might have been a species of "Mixopterus", tentatively designating it as ""Mixopterus" (?) "alveolatus"", Caster and Kjellesvig-Waering assigned the species to "Megalograptus" in 1964, arguing that the morphology of the appendage described by Shuler in 1915 demonstrated that the fossils undoubtedly belonged to "Megalograptus". "M. alveolatus" was kept as a distinct species on account of the third joint of the appendage being proportionally larger than the same joint in "M. ohioensis". In addition  to the five described species assigned to the genus, there may be as many as four distinct undescribed species of "Megalograptus". Caster and Kjellesvig-Waering noted in 1964 that there were very fragmentary eurypterid fossils known from the Katian-age deposits of the Whitewater Formation near Oxford, Ohio, referring these specimens to "Megalograptus". In 2002, fossils belonging to a small variety of "Megalograptus" were first reported from Katian-age deposits of the Nicolet River Formation in Quebec, Canada. "Megalograptus" fossils found in Katian-age deposits in the US state of Georgia and in the Shawangunk Ridge of New York may  also represent two distinct new species. Additionally, fossils potentially referrable to "Megalograptus" have been reported from the Martinsburg Formation of New York and Pennsylvania. Classification. Shortly after being recognized as a eurypterid in the early 20th century, "Megalograptus" was noted by Foerste in 1912 as being similar, and likely closely related, to the genus "Echinognathus". In 1934, Størmer classified "Megalograptus" and "Echinognathus", together with the genera "Mixopterus" and "Carcinosoma", into the family Carcinosomatidae. The taxonomy was amended by Erik N. Kjellesvig-Waering in 1955, who transferred "Mixopterus" to its own family, the Mixopteridae, and placed "Megalograptus" and "Echinognathus" in their  own family, the Megalograptidae. The Megalograptidae has traditionally been interpreted as closely related to the Mixopteridae. In 1964, Caster and Kjellesvig-Waering placed Megalograptidae, alongside Mixopteridae, Carcinosomatidae and Mycteroptidae, into the superfamily Mixopteracea (later renamed to Mixopteroidea). In 1989, Victor P. Tollerton, perceiving them to be distinct enough, placed the Megalograptidae into their own superfamily, the Megalograptoidea. In 2004, O. Erik Tetlie determined "Megalograptus", and by extension the Megalograptidae, to be taxonomically problematic, perceiving the genus to share several potential synapomorphies (derived, "advanced", traits unique to a clade) with both the Eurypteroidea and the Mixopteroidea (now considered a synonym of  Carcinosomatoidea), as well as having a large number of apomorphies (characteristics different from what existed in the ancestor of an organism). Depending on how the analysis was conducted, "Megalograptus" changed position in the phylogenetic tree, only sometimes being recovered as basal within the Mixopteroidea (if taxa where less than one third of the body was preserved were removed). Tetlie speculated that "Megalograptus" and its family could be very basal, given their early age, retaining Megalograptoidea as a distinct superfamily, though with a highly uncertain phylogenetic position, either very basal, between the Onychopterelloidea and the Eurypteroidea, or more derived, between the  Eurypteroidea and the Mixopteroidea. No phylogenetic analysis ever recovered "Megalograptus" in these positions and the genus was often excluded from analyses due to its perceived strange mix of features. The description of "Pentecopterus", the only other genus in the Megalograptidae, in 2015 by Lamsdell and colleagues saw the megalograptids again considered to be close relatives of the mixopterids and carcinosomatids. The phylogenetic analysis accompanying the description of the new genus resolved the Megalograptidae as basal within the relatively derived Carcinosomatoidea superfamily, which also includes the Carcinosomatidae and Mixopteridae. The cladogram below is simplified from the study by Lamsdell "et al ." (2015), collapsed to only display the Carcinosomatoidea. Paleoecology. Discovered alongside specimens of "M. ohioensis" were tube-like structures containing fossil fragments of the trilobite "Isotelus" and of eurypterids, including "M. ohioensis" itself, interpreted as coprolites (fossilized dung). Because of the large size of the coprolites and the presence of fossil material of "M. ohioensis" in and around them, it has been suggested that they are coprolites of "M. ohioensis" itself, thus representing evidence of cannibalism. Because cannibalism is prevalent in modern chelicerates, such as spiders and scorpions (particularly in mating situations), it is possible that eurypterids would have practiced cannibalism  as well. Similar coprolites assigned to the eurypterid "Lanarkopterus dolichoschelus", also from the Ordovician of Ohio, contain, in addition to remains of jawless fish, fragments of smaller specimens of "Lanarkopterus" itself. If the coprolites belong to "Megalograptus", they also further indicate that the genus had a carnivorous diet. The large spines on its forelimbs already indicate that "Megalograptus" was predatory, as they were presumably used for active prey capture, to grasp prey and move it to the mouth. The cercal blades of "Megalograptus" are believed to have been a considerable aid when swimming, acting like a biological rudder, but they  were also able to articulate and move like a scissor. Given that there are no canals for poison in the telson of "Megalograptus", it is possible that they were used almost akin to giant pincers, making the telson and the surrounding structures into a grasping organ, possibly used for defense and during mating. "Megalograptus" is known from what were once near-shore marine environments. "M. ohioensis" occurred alongside a typical Late Ordovician fauna, including trilobites ("Isotelus" and "Flexicalymene"), bryozoans, gastropods, pelecypods, brachiopods, ostracods and scolecodonts. The late Ordovician fossils of "M. ohioensis", as well as the associated fauna, were found  in a rock layer containing remnants of volcanic ash, indicating that the ecosystem in which they lived was destroyed through a volcanic eruption. The fauna co-occurring with "M. shideleri" was similar, consisting of cephalopods, bryozoans, gastropods and scolecodonts. "M. williamsae" was recovered in a so-called "trilobite bed", alongside several different trilobite species. The fossils of "M. welchi" were recovered in a crinoid fossil site otherwise popular with fossil hunters. Other fossil fauna known from the fossil site of "M. welchi" include the trilobites "Ceraurus" and "Dalmanites", the sponge "Brachiospongia" and various crinoids, such as "Glyptocrinus" and "Dendrocrinus". "M.  alveolatus" occurred with a typical Late Ordovician fauna, including brachiopods, such as "Orthorhynchula", and bivalves, such as "Byssonychia".