Type A Genital Appendage

Eurypterids, often informally called sea scorpions, are a group of extinct marine 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) 251.9million 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 ('), meaning 'broad' or 'wide', and ('), meaning 'wing', referring to the pair of wide swimming appendages present in many members of the group.

The eurypterid order includes the largest known arthropods ever to have lived. The largest, ''Jaekelopterus'', reached in length. Eurypterids were not uniformly large and most species were less than long; the smallest eurypterid, '' Alkenopterus'', was only 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.

Description

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). 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.

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. In one lineage, the Pterygotidae, the chelicerae were large and long, with strong, well-developed teeth on specialised chelae (claws). 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.

The opisthosoma comprised 12 segments and the telson, the posteriormost division of the body, which in most species took the form of a blade-like shape. In some lineages, notably the Pterygotioidea, the Hibbertopteridae and the Mycteroptidae, the telson was flattened and may have been used as a rudder while swimming. Some genera within the superfamily Carcinosomatoidea, notably ''Eusarcana'', had a telson similar to that of true scorpions and may even have been capable of using it to inject venom like them. The coxae of the sixth pair of appendages were overlaid by a plate that is referred to as the metastoma, originally derived from a complete exoskeleton segment. The opisthosoma itself can be divided either into a "mesosoma" (comprising segments 1 to 6) and "metasoma" (comprising segments 7 to 12) or into a "preabdomen" (generally comprising segments 1 to 7) and "postabdomen" (generally comprising segments 8 to 12).

The underside of the opisthosoma was covered in structures evolved from modified opisthosomal appendages. Throughout the opisthosoma, these structures formed plate-like structures termed ( in German). These created a branchial chamber (gill tract) between preceding and the ventral surface of the opisthosoma itself, which contained the respiratory organs. The second to sixth opisthosomal segments also contained oval or triangular organs that have been interpreted as organs that aid in respiration. These organs, termed or "gill tracts", would potentially have aided eurypterids to breathe air above water, while , similar to organs in modern horseshoe crabs, would cover the parts that serve for underwater respiration.

The appendages of opisthosomal segments 1 and 2 (the seventh and eighth segments overall) were fused into a structure termed the genital operculum, occupying most of the underside of the opisthosomal segment 2. Near the anterior margin of this structure, the genital appendage (also called the or the median abdominal appendage) protruded. This appendage, often preserved very prominently, has consistently been interpreted as part of the reproductive system and occurs in two recognized types, assumed to correspond to male and female.

Biology

Size

Eurypterids were highly variable in size, depending on factors such as lifestyle, living environment and taxonomic affinity. Sizes around are common in most eurypterid groups. The smallest eurypterid, '' Alkenopterus burglahrensis'', measured just in length.

The largest eurypterid, and the largest known arthropod ever to have lived, is ''Jaekelopterus rhenaniae''. A chelicera from the Emsian Klerf Formation of Willwerath, Germany measured in length, but is missing a quarter of its length, suggesting that the full chelicera would have been long. If the proportions between body length and chelicerae match those of its closest relatives, where the ratio between claw size and body length is relatively consistent, the specimen of ''Jaekelopterus'' that possessed the chelicera in question would have measured between , an average , in length. With the chelicerae extended, another meter (3.28 ft) would be added to this length. This estimate exceeds the maximum body size of all other known giant arthropods by almost half a meter (1.64 ft) even if the extended chelicerae are not included. Two other eurypterids have also been estimated to have reached lengths of 2.5 metres; '' Erettopterus grandis'' (closely related to ''Jaekelopterus'') and '' Hibbertopterus wittebergensis'', but ''E. grandis'' is very fragmentary and the ''H. wittenbergensis'' size estimate is based on trackway evidence, not fossil remains.

The family of ''Jaekelopterus'', the Pterygotidae, is noted for several unusually large species. Both ''Acutiramus'', whose largest member ''A. bohemicus'' measured , and ''Pterygotus,'' whose largest species ''P. grandidentatus'' measured , were gigantic. Several different contributing factors to the large size of the pterygotids have been suggested, including courtship behaviour, predation and competition over environmental resources.

Giant eurypterids were not limited to the family Pterygotidae. An isolated long fossil metastoma of the carcinosomatoid eurypterid '' Carcinosoma punctatum'' indicates the animal would have reached a length of in life, rivalling the pterygotids in size. Another giant was ''Pentecopterus decorahensis'', a primitive carcinosomatoid, which is estimated to have reached lengths of .

Typical of large eurypterids is a lightweight build. Factors such as locomotion, energy costs in molting and respiration, as well as the actual physical properties of the exoskeleton, limit the size that arthropods can reach. A lightweight construction significantly decreases the influence of these factors. Pterygotids were particularly lightweight, with most fossilized large body segments preserving as thin and unmineralized. Lightweight adaptations are present in other giant paleozoic arthropods as well, such as the giant millipede ''Arthropleura'', and are possibly vital for the evolution of giant size in arthropods.

In addition to the lightweight giant eurypterids, some deep-bodied forms in the family Hibbertopteridae were also very large. A carapace from the Carboniferous of Scotland referred to the species ''Hibbertoperus scouleri'' measures wide. As ''Hibbertopterus'' was very wide compared to its length, the animal in question could possibly have measured just short of in length. More robust than the pterygotids, this giant ''Hibbertopterus'' would possibly have rivalled the largest pterygotids in weight, if not surpassed them, and as such be among the heaviest arthropods.

Locomotion

The two eurypterid suborders, Eurypterina and Stylonurina, are distinguished primarily by the morphology of their final pair of appendages. In the Stylonurina, this appendage takes the form of a long and slender walking leg, while in the Eurypterina, the leg is modified and broadened into a swimming paddle. Other than the swimming paddle, the legs of many eurypterines were far too small to do much more than allow them to crawl across the sea floor. In contrast, a number of stylonurines had elongated and powerful legs that might have allowed them to walk on land (similar to modern crabs).

A fossil trackway was discovered in Carboniferous-aged fossil deposits of Scotland in 2005. It was attributed to the stylonurine eurypterid ''Hibbertopterus'' due to a matching size (the trackmaker was estimated to have been about long) and inferred leg anatomy. It is the largest terrestrial trackway—measuring long and averaging in width—made by an arthropod found thus far. It is the first record of land locomotion by a eurypterid. The trackway provides evidence that some eurypterids could survive in terrestrial environments, at least for short periods of time, and reveals information about the stylonurine gait. In ''Hibbertopterus'', as in most eurypterids, the pairs of appendages are different in size (referred to as a heteropodous limb condition). These differently sized pairs would have moved in phase, and the short stride length indicates that ''Hibbertopterus'' crawled with an exceptionally slow speed, at least on land. The large telson was dragged along the ground and left a large central groove behind the animal. Slopes in the tracks at random intervals suggest that the motion was jerky. The gait of smaller stylonurines, such as ''Parastylonurus'', was probably faster and more precise.

The functionality of the eurypterine swimming paddles varied from group to group. In the Eurypteroidea, the paddles were similar in shape to oars. The condition of the joints in their appendages ensured their paddles could only be moved in near-horizontal planes, not upwards or downwards. Some other groups, such as the Pterygotioidea, would not have possessed this condition and were probably able to swim faster. Most eurypterines are generally agreed to have utilized a rowing type of propulsion similar to that of crabs and water beetles. Larger individuals may have been capable of underwater flying (or subaqueous flight) in which the motion and shape of the paddles are enough to generate lift, similar to the swimming of sea turtles and sea lions. This type of movement has a relatively slower acceleration rate than the rowing type, especially since adults have proportionally smaller paddles than juveniles. However, since the larger sizes of adults mean a higher drag coefficient, using this type of propulsion is more energy-efficient.

Some eurypterines, such as ''Mixopterus'' (as inferred from attributed fossil trackways), were not necessarily good swimmers. It likely kept mostly to the bottom, using its swimming paddles for occasional bursts of movements vertically, with the fourth and fifth pairs of appendages positioned backwards to produce minor movement forwards. While walking, it probably used a gait like that of most modern insects. The weight of its long abdomen would have been balanced by two heavy and specialized frontal appendages, and the center of gravity might have been adjustable by raising and positioning the tail.

Preserved fossilized eurypterid trackways tend to be large and heteropodous and often have an associated telson drag mark along the mid-line (as with the Scottish ''Hibbertopterus'' track). Such trackways have been discovered on every continent except for South America. In some places where eurypterid fossil remains are otherwise rare, such as in South Africa and the rest of the former supercontinent Gondwana, the discoveries of trackways both predate and outnumber eurypterid body fossils. Eurypterid trackways have been referred to several ichnogenera, most notably '' Palmichnium'' (defined as a series of four tracks often with an associated drag mark in the mid-line), wherein the holotype of the ichnospecies ''P. kosinkiorum'' preserves the largest eurypterid footprints known to date with the found tracks each being about in diameter. Other eurypterid ichnogenera include ''Merostomichnites'' (though it is likely that many specimens actually represent trackways of crustaceans) and ''Arcuites'' (which preserves grooves made by the swimming appendages).

Respiration

In eurypterids, the respiratory organs were located on the ventral body wall (the underside of the opisthosoma). , evolved from opisthosomal appendages, covered the underside and created a gill chamber where the "gill tracts" were located. Depending on the species, the eurypterid gill tract was either triangular or oval in shape and was possibly raised into a cushion-like state. The surface of this gill tract bore several spinules (small spines), which resulted in an enlarged surface area. It was composed of spongy tissue due to many invaginations in the structure.

Though the is referred to as a "gill tract", it may not necessarily have functioned as actual gills. In other animals, gills are used for oxygen uptake from water and are outgrowths of the body wall. Despite eurypterids clearly being primarily aquatic animals that almost certainly evolved underwater (some eurypterids, such as the pterygotids, would even have been physically unable to walk on land), it is unlikely the "gill tract" contained functional gills when comparing the organ to gills in other invertebrates and even fish. Previous interpretations often identified the eurypterid "gills" as homologous with those of other groups (hence the terminology), with gas exchange occurring within the spongy tract and a pattern of branchio-cardiac and dendritic veins (as in related groups) carrying oxygenated blood into the body. The primary analogy used in previous studies has been horseshoe crabs, though their gill structure and that of eurypterids are remarkably different. In horseshoe crabs, the gills are more complex and composed of many lamellae (plates) which give a larger surface area used for gas exchange. In addition, the gill tract of eurypterids is proportionally much too small to support them if it is analogous to the gills of other groups. To be functional gills, they would have to have been highly efficient and would have required a highly efficient circulatory system. It is considered unlikely, however, that these factors would be enough to explain the large discrepancy between gill tract size and body size.

It has been suggested instead that the "gill tract" was an organ for breathing air, perhaps actually being a lung, plastron or a pseudotrachea. Plastrons are organs that some arthropods evolved secondarily to breathe air underwater. This is considered an unlikely explanation since eurypterids had evolved in water from the start and they would not have organs evolved from air-breathing organs present. In addition, plastrons are generally exposed on outer parts of the body while the eurypterid gill tract is located behind the . Instead, among arthropod respiratory organs, the eurypterid gill tracts most closely resemble the pseudotracheae found in modern isopods. These organs, called pseudotracheae, because of some resemblance to the tracheae (windpipes) of air-breathing organisms, are lung-like and present within the pleopods (back legs) of isopods. The structure of the pseudotracheae has been compared to the spongy structure of the eurypterid gill tracts. It is possible the two organs functioned in the same way.

Some researchers have suggested that eurypterids may have been adapted to an amphibious lifestyle, using the full gill tract structure as gills and the invaginations within it as pseudotrachea. This mode of life may not have been physiologically possible, however, since water pressure would have forced water into the invaginations leading to asphyxiation. Furthermore, most eurypterids would have been aquatic their entire lives. No matter how much time was spent on land, organs for respiration in underwater environments must have been present. True gills, expected to have been located within the branchial chamber within the , remain unknown in eurypterids.

Ontogeny

Like all arthropods, eurypterids matured and grew through static developmental stages referred to as instars. These instars were punctuated by periods during which eurypterids went through ecdysis (molting of the cuticle) after which they underwent rapid and immediate growth. Some arthropods, such as insects and many crustaceans, undergo extreme changes over the course of maturing. Chelicerates, including eurypterids, are in general considered to be direct developers, undergoing no extreme changes after hatching (though extra body segments and extra limbs may be gained over the course of ontogeny in some lineages, such as xiphosurans and sea spiders). Whether eurypterids were true direct developers (with hatchlings more or less being identical to adults) or hemianamorphic direct developers (with extra segments and limbs potentially being added during ontogeny) has been controversial in the past.

Hemianamorphic direct development has been observed in many arthropod groups, such as trilobites, megacheirans, basal crustaceans and basal myriapods. True direct development has on occasion been referred to as a trait unique to arachnids. There have been few studies on eurypterid ontogeny as there is a general lack of specimens in the fossil record that can confidently be stated to represent juveniles. It is possible that many eurypterid species thought to be distinct actually represent juvenile specimens of other species, with paleontologists rarely considering the influence of ontogeny when describing new species.

Studies on a well-preserved fossil assemblage of eurypterids from the Pragian-aged Beartooth Butte Formation

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