Principles of Paleobiology

Reconstruction of the Eurypterid

Pterygotus
By Amo Oliverio, Andrew Kirk, and Janiris Ortiz-Golden

Introduction
The order Eurypterida contains unique specimens of the largest known arthropods. This order of
aquatic chelicerates, commonly called “water scorpions,” existed from the early Late Ordovician to the
Late Permian (Tetlie & Briggs, 2009). There were two significant derived characters among this order,
the development of a large swimming paddle from the posterior-most prosomal appendage and the
enlarged chelicerae. These allowed for chelicerae to be instrumental in the capture of prey (Tetlie
and Briggs, 2009). Clarke & Ruedemann established the family Pterygotidae in 1912 (Miller, 2007).
Pterygotus species, meaning wing-animal or finned one, lived during the Silurian and Lower Devonian
and were among the largest arthropods that ever lived and have reached body lengths of 2m (Tetlie &
Briggs, 2009; Miller, 2007).

Figure 1: Giant arthropods from the fossil record compared with the average height of a human male; (a)
Eurypterid, Early Devonian; (b) Trilobite Isoltelus rex, Late Ordovician; (c) Dragonfly Meganeura monyi, Late
Carboniferous; (d) Millipede Anthropleura armata, Late Carboniferous; Scale bar 50 cm (a—d) (Braddy et al.,
2008)
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The family Pterygotoidea utilized the enlargement of the chelicerae to actively capture prey and
accounts for 56 species (Tetlie & Briggs, 2009). Common characters seen among Pterygotids are:
“enlarged chelicerae, slender non spiniferous appendages III-V, undivided genital appendages, laterally
expanded pretelson and medially carinated, broad paddle-like telson with a complex marginal cuticular
sculpture (Tetlie & Briggs, 2009).” The chelicerae, or “claws,” were likely spinose food-gathering limbs
were thought to capture prey. Then the spineless limbs were used for walking. The broad paddle-like
telson or swimming limb at the posterior completed the appendage plan (Tetlie & Briggs, 2009; Selden,
1984). These Pterygotids can be found in several different sediment types that are rich in magnesium and
calcareous muds (known as “waterlimes”) and shale ranging from the late Early Silurian (428 ma) to the
Middle Devonian (391 ma) (Tetlie 2007). Sediments of this nature are deposited in near-shore marine
locals such as bays, estuaries, and lagoons. These waters are often brackish and are rich ecosystems as
borders between fresh and salt-water habitats. Pterygotids are thought to be the only eurypterids
distributed globally and have been found in rock strata all over the world (Tetlie & Briggs, 2009).

Figure 2: Pterygotus anglicus, ventral view (Nicholson, 1897).

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Figure 3: Pterygotid fossils; (1) Pterygotus (Acutiramus) macrophthalmus cummingsi from the Silurian,
Williamsville waterline at Buffalo, New York. A nearly complete specimen with chelicera reversed, as commonly
found after ecdysis. The joints of the chelicera are indicated. Specimen No. E857 in the Buffalo Museum of
Science. Photography by the Buffalo Museum of Science. X0.5. (2) Same specimen as (1), enlarged to show the
four joints of the left chelicera, in particular the articulation between the first and second joints. X2. Photograph by
the Buffalo Museum of Science. (3) Further enlargement of specimen in (1) to show details of first and second joint
and junction of the two joints. X4. (4) Erettopterus (Truncatiramus) osiliensis (Schmidt), the proximal party of the
chelicera. Silurian, Ludlow Zone K at Kielkond, West Saaremaa (Oesel), Estonia. Specimen No. 7095/5 in the
Museum of Comparative Zoology. X1.8. (5) Erettopterus (Truncatiramus) osiliensis (Schmidt), a free ramus of the
chelicera. Silurian, Ludlow, Zone K at Kielkond, West Saaremaa (Oesel), Estonia. Specimen No. 7059/4 in the
Museum of Comparative Zoology. X1.8. (Kjellesvig-Waering, 1964)

Previous Reconstructions
There are four theories as to the primary habitat for Eurypterids explained by Selden in 1984.
First theory is that eurypterids inhabited freshwater environments and their dead bodies were deposited in
marine habitat. Secondly, eurypterids inhabited primarily freshwater habitats and infrequently visited
marine habitat. Thirdly, they tolerated a wide range of salinities and freely moved or inhabited both fresh
and saline environments. The last theory has a time element, explaining that eurypterids were marine in
the Ordovician, then a wide range of salinities in the Silurian, and gradually became confined to brackish
and fresh-water habitats in the late Paleozoic (Selden, 1984).
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Pterygotid eurypterids are known from North America and have also been found all over the
world. Pterygotus anglicus Agassiz, 1844, from the Lower Devonian of Scotland is perhaps the most
extensively known (Miller 2007). It is believed that most eurypterid remains were the cast exuviae from
molting. The Pterygotids are more abundant in magnesium, calcareous muds and shale. Therefore most
of the Pterygotids are believed to be marine, living near the shore in bays, estuaries and lagoons
(Kjellesvig-Waering, 1964). Specimens of P. anglicus have been found with sharks and placoderms,
demonstrating that part of its life cycle may have been spent in estuaries or lagoons (Miller, 2007)

Originally, the gills of eurypterids were thought to be true gills, making the organisms completely
aquatic (Selden et al, 1985). However, the gills were compared to isopod pseudotracheae and also the
plastron or air-breathing organ of aquatic insects and then the thought was raised that Eurypterids could
have been amphibious (Selden et al, 1985). The area where the true gills would be located is so poorly
fossilized or missing; it is hard to determine what was there or its function, even in Baltoeurpterus
(Selden et al, 1985). Baltoeurpterus was an eurypterid whose natural history is well known and other
eurypterids have gaps in their natural history filled in by comparisons.

Silurian Eurypterids were primary carnivores, and developed armor to protect them from
secondary carnivores (Selden, 1984). No fossilized evidence of diet has been found (Selden,
1984). Large eye organs and comparative studies lead to the belief that Pterygotids had keen
binocular vision (Tetlie & Briggs, 2009). The chelicerae were larger than other genera of
Eurypterids, which improved the ability to catch prey (Tetlie & Briggs, 2009). Also, they had
well defined “teeth” for grasping prey that is trying to escape, which is pictured in Figure 3.
Figure 4 displays the wide range of proposed motion of the chelicerae. It has been debated if they
would have lain on the bottom and ambushed their prey, or actively swam to hunt for food.
 Principles of Paleobiology

Figure 4: Sketches of chelicerae for various species (Kjellesvig-Waering, 1964).

The chelicera were thought to have only three joints, but to have full mobility and to serve as a
means of moving food into the mouth, four articulated joints are necessary, pictured in figure 5. This
would be similar to how crabs feed today. The basal joint would be able to move in almost any direction,
providing full articulation. The first and second joints articulate in a hinge-like fashion and most-likely
lock in a straightened-out position. The third and fourth joints are similarly attached and form the chela.
These chelate are highly developed with teeth. Various suggested functions are associated with these:
defense, sexual display, captures food, and move food to the mouth. (Kjellesvig-Waering, 1964)
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Figure 5: Text-Figures 1-10: (1-10)—Schematic figures representing movements of the four-jointed chelicera of
the Pterygotids; (1-3,8)—Positions for anterior movement; (4)—Twisting action of chelicera; (5,10)—Position
assumed during ecdysis; (6, 7, 9)—Positions during feeding. Should be noted that a triangle must be made before
the chelicerae are in close proximity to the mouth. (Kjellesvig-Waering, 1964).

Individuals in the Family Pterygotidae primarily had streamlined bodies, swimming legs,
marginal eyes, and commonly broad telsons (Selden, 1984). These characteristics provide for a swimming
motion while in aquatic environments. Each of the appendages or limbs served a purpose, not all for
locomotion.
In Pterygotus species, the chelicerae are greatly enlarged for prey capture and are not thought to
be used to assist in locomotion. On their lowest limbs before the tail, swim paddles where developed and
used to provide thrust for swimming. The limbs between the “claws” and the paddles were “walking
limbs”. These were relatively diminutive and thin. They probably enabled the animal to move in a
“buoyant, tripling gait like that of recent lobsters” (Selden, 1984). Pterygotoids are proposed by Selden to
usually swim at low speeds but, had the ability to produce occasional bursts of forward speed using the
flat telson to capture prey.
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Revised Reconstructions

The similarity of the tail-paddle to the large, flat tail fluke of dolphins and whales
indicates the Pterygotus is a powerful swimmer. Finer mobility would be augmented by the
rippling movement of the opisthosoma along the length of the body. Paddle-like front
appendages would have been used for steering as is seen in whales. The lighter and thinner
construction of the carapace adds to the speed and agility of Pterygotus as a swimming predator.
The legs were used for stabilization (Tetlie and Briggs, 2009).

Fossils of fish found among Pterygotus specimens suggest it lived in shallow coastal
areas. The amount and variability of size in groups of fossils found suggest Pterygotus molted.
It is possible that Pterygotus migrated to shore looking for an appropriate site for molting.
During molting, Pterygotus would be left vulnerable to predators and this could have contributed
to its decline. Additionally, it has been suggested that Pterygotus may have become prey to
sharks, as at least one fossil of a shark has been found among Pterygotids. A relatively complete
shark, Doliodus problematicus, was recovered in amongst Pterygotid remains. Armed with
approximately 60 functional teeth, it could have preyed on Pterygotids. (Miller, 2007)

Evidence suggests a life that started in fresh water and moved to a more saline
environment. Air breathing organs are difficult to define (Selden et al, 1985), however, if
Pterygotids were capable of air breathing, and thus terrestrial movement, this would lend towards
an easier method for dispersal and access to more resources. Being that this animal has been
found globally, they must have been a generalist in habit and possibly quite adaptable to many
environments.

The presence of a binocular vision suggests a predatory existence. This is distinctly

different from having eyes on stalks or placed along the sides of a head region as seen in many
other arthropods. Forward facing eyes are essential for predators to produce an overlapping field
of view and create a three dimensional view over as much visual range as possible. The depth of
field is essential to the attack/capture of prey items.

The chelate itself has a simple morphology that functions as a raptorial appendage. The
pivot point between the fixed and free ramus produces an almost linear articulation. This results
in a unique balance between the mechanical advantage and the closing velocity. While the actual
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forces involved in the closing of the chelate cannot be determined, based on the length of the in-
lever arm and out-lever arm provide an estimated mechanical advantage and thus an estimated
velocity ratio. While not a direct correlation, the speed and force the mantis shrimp is capable of
achieving with its raptorial claw demonstrates this potential.

Figure 6: Chelate, Pretelson, and Telson. A, B, C, Pterygotus (Pterygotus); A1, B1, C1, Pterygotus
(Acutiramus); A2, B2, C2, Erettopterus (Erettopterus); A3, B3, C3, Erettopterus (Truncatiramus).
(Kjellesvig-Waering, 1964).

Remains of Pterygotid with chelicerae of different sizes and fish have been noted (Miller, 2007).
Pterygotid eurypterids were probably primary carnivores, feeding on anything smaller than itself that it
could catch in its chelicerae. It is believed that Pterygotids were among the main predators in the middle
Paleozoic world (Miller, 2007). The “toothed” chelicerae probably served the same purpose as a lobster’s
or crawfish’s large powerful claws, which is to capture food and/or defend itself. The slender body and
appendages adapted as “paddles” suggest an animal actively hunting for food. The flattened telson would
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provide a quick burst of speed, either to evade a predator or to aid in capturing prey. However, the
chelicerae seem more adapted for cutting and grasping, than crushing. These would suggest the
possibility of Pterygotus adopting scavenging behavior as a source of food, if needed (Miller, 2007).

Figure 7: The above diagrams were adapted from Selden’s work primarily on Baltoeurpterus (Wikimedia
Commons)

Nevertheless, the posteriorly angled teeth of the chelate with combinations of pointed and
conical teeth are likely used to grasp and spear prey items that are rapidly grabbed by the
extension of the chelicerae. Combining the binocular vision with a fast and agile swimmer and
 Principles of Paleobiology

the presence of chelicerae as deadly tools/weapons the Pterygotus would be a terrifying and
effective predator of the shallow marine environment (Tetlie and Briggs, 2009).

.
Figure 8: Reconstruction of chelicera Pterygotus (Acutiramus) macrophthalmus (Leutze and Heubusch,
1963).

References:

Braddy, Simon J., Markus Poschmann, and O. Erik Tetlie. (2008). "Giant Claw Reveals the Largest Ever
Arthropod." Biology Letters 4.1: 106-09

Ciurca, S.J. and Tetlie O.E. (2007). Society Pterygotids (Chelicerata; Eurypterida) from the
Silurian Vernon Formation of New York. Journal of Paleontology, 81 (4), 725-736.

Kjellesvig-Waering, E. N. (1964). A synopsis of the family Pterygotidae Clarke and Ruedemann,

1912 (Eurypterida). Journal of Paleontology, 331-361.

Leutze, W. P., and C. A. Heubusch. (1963). "Pterygotus in the Bertie Formation (Silurian) of
New York." Journal of Paleontology 37.5: 1080-082.

Miller, R. F. (2007). Pterygotus anglicus Agassiz (Chelicerata: Eurypterida) from Atholville,

Lower Devonian Campbellton Formation, New Brunswick, Canada. Palaeontology, 50(4), 981-
999.

Tetlie, OE. (2007). Distribution and dispersal history of Eurypterida (Chelicerata).

Palaeogeography, Palaeoclimatology, Palaeoecology, 252, 557–574.

Tetlie, O., & Briggs, D. E. (2009). The origin of pterygotid eurypterids (Chelicerata:
Eurypterida). Palaeontology, 52(5), 1141-1148.