Megalograptus

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Megalograptus

Temporal range: Katian, 453–445.2 Ma

PreꞒ

Pg

Replica of the M. ohioensis holotype, Orton

Geology Museum

Scientific classification

Kingdom: Animalia

Phylum: Arthropoda
 Subphylum: Chelicerata

Order: †Eurypterida

Superfamily: †Carcinosomatoidea

Family: †Megalograptidae

Genus: †Megalograptus
Miller, 1874

Type species

†Megalograptus welchi

Miller, 1874

Species

• †M. alveolatus (Shuler,

1915)

• †M. ohioensis Caster &

Kjellesvig-Waering, 1955

• †M. shideleri Caster &

Kjellesvig-Waering, 1964

• †M. welchi Miller, 1874

• †M. williamsae Caster &
Kjellesvig-Waering, 1964

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 78 centimeters (2 ft 7 in).
Some species were substantially smaller, with the smallest, belonging to a hitherto
undetermined species, only growing to about 10 cm (3.9 in) 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.

Contents

• 1Description
o 1.1Size
o 1.2Morphology
o 1.3Coloration
• 2History of research
o 2.1Type material
o 2.2Additional fossils
• 3Classification
• 4Paleoecology
• 5See also
• 6References

Description[edit source]
Size[edit source]
The size of the largest and smallest specimens of M. ohioensis, the largest and best known species
of Megalograptus, compared to a human

Megalograptus was a genus of large megalograptid eurypterids. The largest species

was M. ohioensis, which ranged in length from 49 to 78 centimeters (1 foot 7 inches
to 2 feet 7 inches). 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 170 cm (5 ft 7 in).[1] 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 150 cm (4 ft
11 in) 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.[2] In the case of M. shideleri, once estimated to have reached lengths of
200 cm (6 ft 7 in),[2] 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 56 cm (1 ft
10 in).[1] The length of the species M. alveolatus is uncertain,[2] but it was much
smaller than M. ohioensis,[3] and M. williamsae grew to about 50 cm (1 ft 8 in). The
smallest known species of Megalograptus was an as yet undescribed Canadian
species which only grew to 10 cm (3.9 in) in length.[2]
Morphology[edit source]
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. [3]
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.[3]

Anatomical diagram of the upper side of M. ohioensis, featuring some of the body parts mentioned in the
text

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).[3]
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.[3] Cercal
blades are only known from one other eurypterid, Holmipterus,[4] and are lacking in
the basal ("primitive") megalograptid Pentecopterus.[1] 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.[3]
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.[3]
Coloration[edit source]

M. ohioensis, depicted with bristles and its inferred dark brown and black life coloration, per its 1964
description by Caster and Kjellesvig-Waering.

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.[3] A similar, but darker, brown and black color scheme has been inferred for M.
ohioensis,[5] 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. [3] 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. [5]
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. [3]

History of research[edit source]

Type material[edit source]

The M. welchi type material; a "walking leg" (top), a fragmentary "walking leg" (bottom right) and a
postabdominal segment (bottom left) (left) and the same fossils illustrated by Miller in 1874 as remains of
a graptolite, covered in "polyp cells" (right)

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),[3] and gave it the name Megalograptus, meaning "great
writing" (deriving from the Greek megale, "great", and graptos, "writing", commonly
used for graptolite fossils).[6] One reason for Miller's mistaken identification is that the
exact outline of the fossils was unclear because they were not properly cleaned yet. [7]
The fragmentary fossils of M. welchi were initially recovered by L. B. Welch, whom
the species name welchi honours, near Liberty, Ohio,[3] in rocks of Katian (Late
Ordovician) age[2] of the Elkhorn Formation.[3] 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.[3][7]
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.[3] 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.[3][7] 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.[3] 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.[7]
Additional fossils[edit source]

The fragmentary fossil type material of M. alveolatus, originally assigned to the genus Ctenopterus

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.[3] These fossils, found in
deposits of Katian age[2] 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).[3]
In 1964, Caster and Kjellesvig-Waering named two new species
of Megalograptus[3] based on additional fossil material from the Katian [2] 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.[3]
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, [8] collected in Late Ordovician[2] deposits
along Walker Mountain in Virginia[8] belonging to the Bays Formation.[9] It was the first
eurypterid to be described from Virginia.[10] The species name alveolatus refers to the
pronounced development of the alveolar processes (pits) around the
spines.[8] 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",[10] 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.[3]
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 [3] known
from the Katian-age[2] deposits of the Whitewater Formation near Oxford, Ohio,
referring these specimens to Megalograptus.[3] 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.[2][11][12] 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.[2][12] Additionally, fossils potentially referrable
to Megalograptus have been reported from the Martinsburg Formation of New York
and Pennsylvania.[13]

Classification[edit source]
M. ohioensis (left) compared to the earliest and basalmost known member of
the Megalograptidae, Pentecopterus decorahensis (right)

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.[7] 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.[14] The Megalograptidae has traditionally been interpreted as closely
related to the Mixopteridae.[15] In 1964, Caster and Kjellesvig-Waering placed
Megalograptidae, alongside Mixopteridae, Carcinosomatidae and Mycteroptidae, into
the superfamily Mixopteracea (later renamed to Mixopteroidea). [3] In 1989, Victor P.
Tollerton, perceiving them to be distinct enough, placed the Megalograptidae into
their own superfamily, the Megalograptoidea. [16]
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,[15] 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.[12] 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. [1]
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.[1]
Carcinosomatoidea   Megalograptidae  
    Pentecopterus decorahensis
   
   
Echinognathus clevelandi
   
   
Megalograptus williamsae
   
   
Megalograptus ohioensis
   
 
Megalograptus welchi
 

  Mixopteridae  
  Lanarkopterus dolichoschelus
   
   
Mixopterus kiaeri
   
 
Mixopterus multispinosus
 
Carcinosomatidae  
Holmipterus suecicus
   
 
Rhinocarcinosoma vaningeni
 
     
      Carcinosoma newlini
 
  
Carcinosoma libertyi

   
Eusarcana acrocephalus
   
 
Eusarcana scorpionis
 

Paleoecology[edit source]

Simplified reconstruction of the telson-pretelson assemblage of M. ohioensis, possibly used for grasping,
viewed from below (left) and above (right)

Discovered alongside specimens of M. ohioensis were tube-like structures containing

fossil fragments of the trilobite Isotelus and of eurypterids, including M.
ohioensis itself,[3] interpreted as coprolites (fossilized dung).[3][17] 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.[3] 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.[17] If the coprolites belong
to Megalograptus, they also further indicate that the genus had a carnivorous
diet.[3] The large spines on its forelimbs already indicate that Megalograptus was
predatory,[3] as they were presumably used for active prey capture, [18] to grasp prey
and move it to the mouth.[3] The cercal blades of Megalograptus are believed to have
been a considerable aid when swimming, acting like a biological rudder,[1][3] 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. [3]
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, ostra
cods 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.[3] 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.[7] M. alveolatus occurred with a typical Late
Ordovician fauna,[3] including brachiopods, such as Orthorhynchula, and bivalves,
such as Byssonychia.[19]

See also[edit source]

• List of eurypterid genera
• Timeline of eurypterid research

References[edit source]
1. ^ Jump up to:a b c d e f Lamsdell, James C.; Briggs, Derek E. G.; Liu, Huaibao P.; Witzke,
Brian J.; McKay, Robert M. (2015). "The oldest described eurypterid: a giant Middle
Ordovician (Darriwilian) megalograptid from the Winneshiek Lagerstätte of Iowa". BMC
Evolutionary Biology. 15: 169. doi:10.1186/s12862-015-0443-9. ISSN 1471-
2148. PMC 4556007. PMID 26324341.
2. ^ Jump up to:a b c d e f g h i j k Lamsdell, James C.; Braddy, Simon J. (2009). "Cope's rule
and Romer's theory: patterns of diversity and gigantism in eurypterids and Palaeozoic
vertebrates". Biology Letters. 6 (2): 265–
269. doi:10.1098/rsbl.2009.0700. PMC 2865068. PMID 19828493. Supplementary
information
3. ^ Jump up to:a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah aiCaster, Kenneth E.;
Kjellesvig-Waering, Erik N. (1964). "Upper Ordovician eurypterids of
Ohio". Paleontological Research Institution. 4: 301–342.
4. ^ Kjellesvig-Waering, Erik N. (1979). "Eurypterids" (PDF). In Laufeld, Sven; Skoglund,
Roland (eds.). Lower Wenlock faunal and floral dynamics – Vattenfallet section,
Gotland (PDF). Sveriges geologiska undersökning: Avhandlingar och uppsatser. pp. 121–
136. ISBN 978-9-171-58170-9.
5. ^ Jump up to:a b Kjellesvig-Waering, Erik N. (1958). "Some Previously Unknown
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