Eurypterid


Eurypterids, often informally called sea scorpions, are a group of extinct arthropods that form the order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period 467.3 million years ago. The group is likely to have appeared first either during the Early Ordovician or Late Cambrian period. With approximately 250 species, the Eurypterida is the most diverse Paleozoic chelicerate order. Following their appearance during the Ordovician, eurypterids became major components of marine faunas during the Silurian, from which the majority of eurypterid species have been described. The Silurian genus Eurypterus accounts for more than 90% of all known eurypterid specimens. Though the group continued to diversify during the subsequent Devonian period, the eurypterids were heavily affected by the Late Devonian extinction event. They declined in numbers and diversity until becoming extinct during the Permian–Triassic extinction event (or sometime shortly before) 251.9 million years ago.

Although popularly called "sea scorpions", only the earliest eurypterids were marine; many later forms lived in brackish or fresh water, and they were not true scorpions. Some studies suggest that a dual respiratory system was present, which would have allowed for short periods of time in terrestrial environments. The name Eurypterida comes from the Ancient Greek words εὐρύς (eurús), meaning 'broad' or 'wide', and πτερόν (pterón), meaning 'wing', referring to the pair of wide swimming appendages present in many members of the group.

The eurypterids include the largest known arthropods ever to have lived. The largest, Jaekelopterus, reached 2.5 meters (8.2 ft) in length. Eurypterids were not uniformly large and most species were less than 20 centimeters (8 in) long; the smallest eurypterid, Alkenopterus, was only 2.03 centimeters (0.80 in) long. Eurypterid fossils have been recovered from every continent. A majority of fossils are from fossil sites in North America and Europe because the group lived primarily in the waters around and within the ancient supercontinent of Euramerica. Only a handful of eurypterid groups spread beyond the confines of Euramerica and a few genera, such as Adelophthalmus and Pterygotus, achieved a cosmopolitan distribution with fossils being found worldwide.

Like all other arthropods, eurypterids possessed segmented bodies and jointed appendages (limbs) covered in a cuticle composed of proteins and chitin. As in other chelicerates, the body was divided into two tagmata (sections); the frontal prosoma (head) and posterior opisthosoma (abdomen).[1] The prosoma was covered by a carapace (sometimes called the "prosomal shield") on which both compound eyes and the ocelli (simple eye-like sensory organs) were located.[2]

The prosoma also bore six pairs of appendages which are usually referred to as appendage pairs I to VI. The first pair of appendages, the only pair placed before the mouth, is called the chelicerae (homologous to the fangs of spiders). They were equipped with small pincers used to manipulate food fragments and push them into the mouth.[2] In one lineage, the Pterygotidae, the chelicerae were large and long, with strong, well-developed teeth on specialised chelae (claws).[3] The subsequent pairs of appendages, numbers II to VI, possessed gnathobases (or "tooth-plates") on the coxae (limb segments) used for feeding. These appendages were generally walking legs that were cylindrical in shape and were covered in spines in some species. In most lineages, the limbs tended to get larger the farther back they were. In the Eurypterina suborder, the larger of the two eurypterid suborders, the sixth pair of appendages was also modified into a swimming paddle to aid in traversing aquatic environments.[2]


Restoration of Eurypterus with body parts labelled
Size comparison of six of the largest eurypterids: Pterygotus grandidentatus, Pentecopterus decorahensis, Acutiramus macrophthalmus, A. bohemicus, Carcinosoma punctatum, and Jaekelopterus rhenaniae
Illustration of Hibbertopterus, a large stylonurine (lacking swimming paddles) eurypterid
The holotype of Palmichnium kosinkiorum, containing the largest eurypterid footprints known
The supposed "gill tracts" of eurypterids have been compared to the air-breathing pseudotracheae present in the posterior legs of modern isopods, such as Oniscus (pictured).
Larval (left) and juvenile (right) instars of Strobilopterus (not to scale)
Pterygotus depicted hunting Birkenia
Type A genital appendage of Adelophthalmus mansfieldi
Type B genital appendage of Kokomopterus longicaudatus
Type A genital appendage of Eurypterus sp.
A reconstruction of Pentecopterus, the earliest known eurypterid. The family to which Pentecopterus belongs, the Megalograptidae, was the first truly successful eurypterid group.
Reconstruction of Erettopterus, a member of the highly successful Silurian and Devonian eurypterid family Pterygotidae
Reconstruction of Adelophthalmus, the only eurypterine (with swimming paddles) eurypterid to survive the Late Devonian extinction and persist into the subsequent Carboniferous and Permian periods.
Reconstruction of the hibbertopterid Campylocephalus, the last known surviving eurypterid. Campylocephalus went extinct during the Permian-Triassic extinction event or sometime shortly before.
Figure of Eurypterus remipes by James E. De Kay (1825).
Evolutionary tree of eurypterids as imagined by John Mason Clarke and Rudolf Ruedemann in 1912.
Eurypterids have historically been seen as closely related to xiphosurans such as the Atlantic horseshoe crab (top), united in the class Merostomata. Recent studies are in favor of a closer relationship to arachnids such as Heterophrynus (bottom), united under the clade Sclerophorata.
The most important taxonomic character in eurypterids is the morphology of the sixth pair of appendages. In most eurypterids of the suborder Eurypterina, such as Bassipterus (top), this leg is modified into a swimming paddle. In eurypterids of the suborder Stylonurina, such as Parastylonurus (bottom), it is not.