Eurypterid
Eurypterid | |
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Fossil specimen of | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Subphylum: | Chelicerata |
Clade: | Sclerophorata
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Order: | †Eurypterida Burmeister, 1843 |
Suborders | |
Incertae sedis | |
Synonyms | |
Eurypterids, often informally called sea scorpions, are a group of extinct
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 eurypterid order includes the largest known arthropods ever to have lived. The largest,
Like all other
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
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.[2] 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 modern scorpions and may have been capable of using it to inject venom.[4][5] 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).[2]
The underside of the opisthosoma was covered in structures evolved from modified opisthosomal appendages. Throughout the opisthosoma, these structures formed plate-like structures termed Blattfüsse (lit. 'leaf-feet' in German).
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
Biology
Size
Eurypterids were highly variable in size, depending on factors such as lifestyle, living environment and taxonomic affinity. Sizes around 100 centimeters (3.3 ft) are common in most eurypterid groups.[7] The smallest eurypterid, Alkenopterus burglahrensis, measured just 2.03 centimeters (0.80 in) in length.[8]
The largest eurypterid, and the largest known arthropod ever to have lived, is
The family of Jaekelopterus, the Pterygotidae, is noted for several unusually large species. Both Acutiramus, whose largest member A. bohemicus measured 2.1 meters (6.9 ft), and Pterygotus, whose largest species P. grandidentatus measured 1.75 meters (5.7 ft), were gigantic.[9] Several different contributing factors to the large size of the pterygotids have been suggested, including courtship behaviour, predation and competition over environmental resources.[11]
Giant eurypterids were not limited to the family Pterygotidae. An isolated 12.7 centimeters (5.0 in) long fossil metastoma of the carcinosomatoid eurypterid Carcinosoma punctatum indicates the animal would have reached a length of 2.2 meters (7.2 ft) in life, rivalling the pterygotids in size.[12] Another giant was Pentecopterus decorahensis, a primitive carcinosomatoid, which is estimated to have reached lengths of 1.7 meters (5.6 ft).[13]
Typical of large eurypterids is a lightweight build. Factors such as locomotion, energy costs in
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 65 cm (26 in) wide. As Hibbertopterus was very wide compared to its length, the animal in question could possibly have measured just short of 2 meters (6.6 ft) 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.[15]
Locomotion
The two eurypterid suborders, Eurypterina and
A
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.[20] 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.[21]
Some eurypterines, such as
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.[23] 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),[24] wherein the holotype of the ichnospecies P. kosinkiorum preserves the largest eurypterid footprints known to date with the found tracks each being about 7.6 centimeters (3.0 in) in diameter.[25] 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).[24][26][27]
Respiration
In eurypterids, the respiratory organs were located on the ventral body wall (the underside of the opisthosoma). Blattfüsse, evolved from opisthosomal appendages, covered the underside and created a gill chamber where the "gill tracts" were located.[2] 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.[28]
Though the Kiemenplatte 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.[29]
It has been suggested instead that the "gill tract" was an organ for breathing air, perhaps actually being a
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
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.[33]
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.[33] 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.[34]
Studies on a well-preserved fossil assemblage of eurypterids from the Pragian-aged Beartooth Butte Formation in Cottonwood Canyon, Wyoming, composed of multiple specimens of various developmental stages of eurypterids Jaekelopterus and Strobilopterus, revealed that eurypterid ontogeny was more or less parallel and similar to that of extinct and extant xiphosurans, with the largest exception being that eurypterids hatched with a full set of appendages and opisthosomal segments. Eurypterids were thus not hemianamorphic direct developers, but true direct developers like modern arachnids.[35]
The most frequently observed change occurring through ontogeny (except for some genera, such as Eurypterus, which appear to have been static) is the metastoma becoming proportionally less wide. This ontogenetic change has been observed in members of several superfamilies, such as the Eurypteroidea, the Pterygotioidea and the
Feeding
No fossil gut contents from eurypterids are known, so direct evidence of their diet is lacking. The eurypterid biology is particularly suggestive of a carnivorous lifestyle. Not only were many large (in general, most predators tend to be larger than their prey), but they had
Fossils preserving
Eurypterid coprolites discovered in deposits of Ordovician age in Ohio containing fragments of a trilobite and eurypterid Megalograptus ohioensis in association with full specimens of the same eurypterid species have been suggested to represent evidence of
Though apex predatory roles would have been limited to the very largest eurypterids, smaller eurypterids were likely formidable predators in their own right just like their larger relatives.[7]
Reproductive biology
As in many other entirely extinct groups, understanding and researching the reproduction and sexual dimorphism of eurypterids is difficult, as they are only known from fossilized shells and carapaces. In some cases, there might not be enough apparent differences to separate the sexes based on morphology alone.[17] Sometimes two sexes of the same species have been interpreted as two different species, as was the case with two species of Drepanopterus (D. bembycoides and D. lobatus).[40]
The eurypterid prosoma is made up of the first six exoskeleton segments fused together into a larger structure. The seventh segment (thus the first opisthosomal segment) is referred to as the metastoma and the eighth segment (distinctly plate-like) is called the operculum and contains the genital aperature. The underside of this segment is occupied by the genital operculum, a structure originally evolved from ancestral seventh and eighth pair of appendages. In its center, as in modern horseshoe crabs, is a genital appendage. This appendage, an elongated rod with an internal duct, is found in two distinct morphs, generally referred to as "type A" and "type B".[17] These genital appendages are often preserved prominently in fossils and have been the subject of various interpretations of eurypterid reproduction and sexual dimorphism.[41]
Type A appendages are generally longer than those of type B. In some genera they are divided into different numbers of sections, such as in Eurypterus where the type A appendage is divided into three but the type B appendage into only two.
The primary function of the long, assumed female, type A appendages was likely to take up spermatophore from the substrate into the
Type B appendages, assumed male, would have produced, stored and perhaps shaped spermatophore in a heart-shaped structure on the dorsal surface of the appendage. A broad genital opening would have allowed large amounts of spermatophore to be released at once. The long furca associated with type B appendages, perhaps capable of being lowered like the type A appendage, could have been used to detect whether a substrate was suitable for spermatophore deposition.[48]
Evolutionary history
Origins
Until 1882 no eurypterids were known from before the Silurian. Contemporary discoveries since the 1880s have expanded the knowledge of early eurypterids from the
Pentecopterus was a relatively derived eurypterid, part of the megalograptid family within the carcinosomatoid superfamily. Its derived position suggests that most eurypterid clades, at least within the eurypterine suborder, had already been established at this point during the Middle Ordovician.[52] The earliest known stylonurine eurypterid, Brachyopterus,[7] is also Middle Ordovician in age. The presence of members of both suborders indicates that primitive stem-eurypterids would have preceded them, though these are so far unknown in the fossil record. The presence of several eurypterid clades during the Middle Ordovician suggests that eurypterids either originated during the Early Ordovician and experienced a rapid and explosive radiation and diversification soon after the first forms evolved, or that the group originated much earlier, perhaps during the Cambrian period.[52]
As such, the exact eurypterid time of origin remains unknown. Though fossils referred to as "primitive eurypterids" have occasionally been described from deposits of Cambrian or even
The fossil record of Ordovician eurypterids is quite poor. The majority of eurypterids once reportedly known from the Ordovician have since proven to be misidentifications or
Silurian
Eurypterids were most diverse and abundant between the Middle Silurian and the Early Devonian, with an absolute peak in diversity during the
Though stylonurine eurypterids generally remained rare and low in number, as had been the case during the preceding Ordovician, eurypterine eurypterids experienced a rapid rise in diversity and number.[57] In most Silurian fossil beds, eurypterine eurypterids account for 90% of all eurypterids present.[58] Though some were likely already present by the Late Ordovician (simply missing from the fossil record so far),[52] a vast majority of eurypterid groups are first recorded in strata of Silurian age. These include both stylonurine groups such as the Stylonuroidea, Kokomopteroidea and Mycteropoidea as well as eurypterine groups such as the Pterygotioidea, Eurypteroidea and Waeringopteroidea.[59]
The most successful eurypterid by far was the Middle to Late Silurian Eurypterus, a
During the Late Silurian the pterygotid eurypterids, large and specialized forms with several new adaptations, such as large and flattened telsons capable of being used as rudders, and large and specialized chelicerae with enlarged pincers for handling (and potentially in some cases killing) prey appeared.[3][4] Though the largest members of the family appeared in the Devonian, large two meter (6.5+ ft) pterygotids such as Acutiramus were already present during the Late Silurian.[10] Their ecology ranged from generalized predatory behavior to ambush predation and some, such as Pterygotus itself, were active apex predators in Late Silurian marine ecosystems.[63] The pterygotids were also evidently capable of crossing oceans, becoming one of only two eurypterid groups to achieve a cosmopolitan distribution.[64]
Devonian
Though the eurypterids continued to be abundant and diversify during the Early Devonian (for instance leading to the evolution of the pterygotid Jaekelopterus, the largest of all arthropods), the group was one of many heavily affected by the Late Devonian extinction. The extinction event, only known to affect marine life (particularly trilobites, brachiopods and reef-building organisms) effectively crippled the abundance and diversity previously seen within the eurypterids.[65]
A major decline in diversity had already begun during the Early Devonian and eurypterids were rare in marine environments by the Late Devonian. During the Frasnian stage four families went extinct, and the later Famennian saw an additional five families going extinct.[65] As marine groups were the most affected, the eurypterids were primarily impacted within the eurypterine suborder. Only one group of stylonurines (the family Parastylonuridae) went extinct in the Early Devonian. Only two families of eurypterines survived into the Late Devonian at all (Adelophthalmidae and Waeringopteridae). The eurypterines experienced their most major declines in the Early Devonian, during which over 50% of their diversity was lost in just 10 million years. Stylonurines, on the other hand, persisted through the period with more or less consistent diversity and abundance but were affected during the Late Devonian, when many of the older groups were replaced by new forms in the families Mycteroptidae and Hibbertopteridae.[66]
It is possible that the catastrophic extinction patterns seen in the eurypterine suborder were related to the emergence of more derived fish. Eurypterine decline began at the point when jawless fish first became more developed and coincides with the emergence of
Stylonurines of the surviving hibbertopterid and mycteroptid families completely avoided competition with fish by evolving towards a new and distinct ecological niche. These families experienced a radiation and diversification through the Late Devonian and Early Carboniferous, the last ever radiation within the eurypterids, which gave rise to several new forms capable of "sweep-feeding" (raking through the substrate in search of prey).[68]
Carboniferous and Permian
Only three eurypterid families—Adelophthalmidae, Hibbertopteridae and Mycteroptidae—survived the extinction event in its entirety. We used to think that these were all freshwater animals, which would have rendered the eurypterids extinct in marine environments.[65], and with marine eurypterid predators gone, sarcopterygians, such as the rhizodonts, were the new apex predators in marine environments.[67] However, various recent findings raise doubts about this, and suggest that these eurypterids were euryhaline forms that lived in marginal marine environments, such as estuaries, deltas, lagoons, and coastal ponds. One argument is paleobiogeographical; pterygotoid distribution seems to require oceanic dispersal.[69] A recent review of Adelophthalmoidea admitted that "There is much more of a marine influence in many of the sections yielding Adelophthalmus than has previously been acknowledged."[70] Similarly, a study of the eurypterid Hibbertopterus from the Carboniferous of New Mexico concluded that the habitat of some eurypterids "may need to be re-evaluated".[71] The sole surviving eurypterine family, Adelophthalmidae, was represented by only a single genus, Adelophthalmus. The hibbertopterids, mycteroptids and Adelophthalmus survived into the Permian.[72]
Adelophthalmus became the most common of all late Paleozoic eurypterids, existing in greater number and diversity than surviving stylonurines, and diversified in the absence of other eurypterines.[73] Out of the 33 species referred to Adelophthalmus, 23 (69%) are from the Carboniferous alone.[74][75] The genus reached its peak diversity in the Late Carboniferous. Though Adelophthalmus had already been relatively widespread and represented around all major landmasses in the Late Devonian, the amalgamation of Pangaea into a global supercontinent over the course of the last two periods of the Paleozoic allowed Adelophthalmus to gain an almost worldwide distribution.[58]
During the Late Carboniferous and
Mycteroptids and hibbertopterids continued to survive for some time, with one genus of each group known from Permian strata: Hastimima and Campylocephalus respectively.[77] Hastimima went extinct during the Early Permian,[78] as Adelophthalmus had, while Campylocephalus persisted longer. A massive incomplete carapace from Permian deposits in Russia represents the sole fossil remains of the species C. permianus, which might have reached 1.4 meters (4.6 ft) in length, while originally considered to be Late Permian in age[10] other sources suggest a Kungurian-Roadian age for the specimen.[79] A fragment of eurypterid cuticle, given the species name Woodwardopterus freemanorum is known from the terminal Permian of Australia, which represents the youngest known eurypterid.[79] No eurypterids are known from fossil beds higher than the Permian. This indicates that the last eurypterids died either in the catastrophic extinction event at its end or at some point shortly before it. This extinction event, the Permian–Triassic extinction event, is the most devastating mass extinction recorded, and rendered many other successful Paleozoic groups, such as the trilobites, extinct.[80]
History of study
The first known eurypterid specimen was discovered in the Silurian-aged rocks of New York, to this day one of the richest eurypterid fossil locations. Samuel L. Mitchill described the specimen, discovered near Westmoreland in Oneida county in 1818. He erroneously identified the fossil as an example of the fish Silurus, likely due to the strange, catfish-like appearance of the carapace. Seven years later, in 1825, James E. DeKay examined the fossil and recognized it as clearly belonging to an arthropod. He thought the fossil, which he named Eurypterus remipes, represented a crustacean of the order Branchiopoda, and suggested it might represent a missing link between the trilobites and more derived branchiopods.[81] The name Eurypterus derives from Greek eurús (εὐρύς 'broad, wide') and pteron (πτερόν 'wing').[82]
In 1843, Hermann Burmeister published his view on trilobite taxonomy and how the group related to other organisms, living and extinct, in the work Die Organisation der Trilobiten aus ihren lebenden Verwandten entwickelt. He considered the trilobites to be crustaceans, as previous authors had, and classified them together with what he assumed to be their closest relatives, Eurypterus and the genus Cytherina, within a clade he named "Palaeadae". Within Palaeadae, Burmeister erected three families; the "Trilobitae" (composed of all trilobites), the "Cytherinidae" (composed only of Cytherina, an animal today seen as an ostracod) and the Eurypteridae (composed of Eurypterus, then including three species).[83]
The fourth eurypterid genus to be described (following Hibbertopterus in 1836 and Campylocephalus in 1838, not identified as eurypterids until later), out of those still seen as taxonomically valid in modern times, was Pterygotus (lit. 'winged one'), described by Louis Agassiz in 1839.[84] Pterygotus was considerably larger in size than Eurypterus and when the first fossils were discovered by quarrymen in Scotland they were referred to as "Seraphims" by the workers. Agassiz first thought the fossils represented remains of fish, only recognizing their nature as arthropod remains five years later in 1844.[85]
In 1849, Frederick M'Coy classified Pterygotus together with Eurypterus and Belinurus (a genus today seen as a xiphosuran) within Burmeister's Eurypteridae. M'Coy considered the Eurypteridae to be a group of crustaceans within the order Entomostraca, closely related to horseshoe crabs.[86] A fourth genus, Slimonia, based on fossil remains previously assigned to a new species of Pterygotus, was referred to the Eurypteridae in 1856 by David Page.[87]
Jan Nieszkowski's De Euryptero Remipede (1858) featured an extensive description of Eurypterus fischeri (now seen as synonymous with another species of Eurypterus, E. tetragonophthalmus), which, along with the monograph On the Genus Pterygotus by Thomas Henry Huxley and John William Salter, and an exhaustive description of the various eurypterids of New York in Volume 3 of the Palaeontology of New York (1859) by James Hall, contributed massively to the understanding of eurypterid diversity and biology. These publications were the first to fully describe the whole anatomy of eurypterids, recognizing the full number of prosomal appendages and the number of preabdominal and postabdominal segments. Both Nieszkowski and Hall recognized that the eurypterids were closely related to modern chelicerates, such as horseshoe crabs.[88]
In 1865, Henry Woodward described the genus Stylonurus (named and figured, but not thoroughly described, by David Page in 1856) and raised the rank of the Eurypteridae to that of order, effectively creating the Eurypterida as the taxonomic unit it is seen as today.[89] In the work Anatomy and Relations of the Eurypterida (1893), Malcolm Laurie added considerably to the knowledge and discussion of eurypterid anatomy and relations. He focused on how the eurypterids related to each other and to trilobites, crustaceans, scorpions, other arachnids and horseshoe crabs. The description of Eurypterus fischeri by Gerhard Holm in 1896 was so elaborate that the species became one of the most completely known of all extinct animals, so much so that the knowledge of E. fischeri was comparable with the knowledge of its modern relatives (such as the Atlantic horseshoe crab). The description also helped solidify the close relationship between the eurypterids and other chelicerates by showcasing numerous homologies between the two groups.[90]
In 1912,
Classification
Historically, a close relationship between eurypterids and xiphosurans (such as the modern Atlantic horseshoe crab) has been assumed by most researchers. Several homologies encourage this view, such as correlating segments of the appendages and the prosoma. Additionally, the presence of plate-like appendages bearing the "gill tracts" on appendages of the opisthosoma (the Blattfüssen) was cited early as an important homology. In the last few decades of the nineteenth century, further homologies were established, such as the similar structures of the compound eyes of Pterygotus and horseshoe crabs (seen as especially decisive as the eye of the horseshoe crab was seen as possessing an almost unique structure) and similarities in the ontogeny within both groups.
Due to these similarities, the xiphosurans and eurypterids have often been united under a single class or
In 1866, Ernst Haeckel classified the Merostomata (containing virtually only the Eurypterida) and Xiphosura within a group he named Gigantostraca within the crustaceans. Though Haeckel did not designate any taxonomic rank for this clade, it was interpreted as equivalent to the rank of subclass, such as the Malacostraca and Entomostraca, by later researchers such as John Sterling Kinsgsley.[96] In subsequent research, Gigantostraca has been treated as synonymous with Merostomata (rarely) and Eurypterida itself (more commonly).[97][98]
A phylogenetic analysis (the results presented in a cladogram below) conducted by James Lamsdell in 2013 on the relationships within the Xiphosura and the relations to other closely related groups (including the eurypterids, which were represented in the analysis by genera Eurypterus, Parastylonurus,
Arthropoda
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Internal relationships
The internal classification of eurypterids within the Eurypterida is based mainly on eleven established characters. These have been used throughout the history of eurypterid research to establish clades and genera. These characters include: the shape of the prosoma, the shape of the metastoma, the shape and position of the eyes, the types of prosomal appendages, the types of swimming leg paddles, the structure of the doublure (the fringe of the dorsal exoskeleton), the structure of the opithosoma, the structure of the genital appendages, the shape of the telson and the type of ornamentation present. It is worth noting that not all of these characters are of equal taxonomic importance.[101] They are not applicable to all eurypterids either; stylonurine eurypterids lack swimming leg paddles entirely.[16] Some characters, including the prosoma and metastoma shapes and the position and shapes of the eyes, are seen as important only for the distinction between different genera.[102] Most superfamilies and families are defined based on the morphology of the appendages.[103]
The most important character used in eurypterid taxonomy is the type of prosomal appendages as this character is used to define entire suborders. General leg anatomy can also be used to define superfamilies and families. Historically, the chelicerae were considered the most important appendages from a taxonomical standpoint since they only occurred in two general types: a eurypterid type with small and toothless pincers and a pterygotid type with large pincers and teeth. This distinction has historically been used to divide the Eurypterida into the two suborders Eurypterina (small chelicerae) and "Pterygotina" (large and powerful chelicerae).[104] This classification scheme is not without problems. In Victor Tollerton's 1989 taxonomic revision of the Eurypterida, with suborders Eurypterina and Pterygotina recognized, several clades of eurypterids today recognized as stylonurines (including hibbertopterids and mycteroptids) were reclassified as non-eurypterids in the new separate order "Cyrtoctenida" on the grounds of perceived inconsistencies in the prosomal appendages.[105]
Modern research favors a classification into suborders Eurypterina and Stylonurina instead, supported by phylogenetic analyses.[106][36] In particular, pterygotid eurypterids share a number of homologies with derived eurypterine eurypterids such as the adelophthalmids, and are thus best classified as derived members of the same suborder.[107] In the Stylonurina, the sixth pair of appendages is represented by long and slender walking legs and lack a modified spine (referred to as the podomere 7a). In most eurypterids in the Eurypterina, the sixth pair of appendages is broadened into swimming paddles and always has a podomere 7a. 75% of eurypterid species are eurypterines and they represent 99% of all fossil eurypterid specimens.[16] Of all eurypterid clades, the Pterygotioidea is the most species-rich, with over 50 species. The second most species-rich clade is the Adelophthalmoidea, with over 40 species.[58]
The cladogram presented below, covering all currently recognized eurypterid families, follows a 2007 study by O. Erik Tetlie.
Eurypterida | |
See also
- List of eurypterid genera
- Nepidae—an unrelated family of insects, commonly known as "water scorpions".
- Cottidae—family of fishes in which some members contain "sea scorpion" in their common name.
- History of life
References
Citations
- ^ Størmer 1955, p. 23.
- ^ a b c d e f g h i Braddy & Dunlop 1997, pp. 437–439.
- ^ a b Tetlie & Briggs 2009, p. 1141.
- ^ a b Plotnick & Baumiller 1988, p. 22.
- ^ Clarke & Ruedemann 1912, p. 244.
- ^ "Selectivity in the evolution of Palaeozoic arthropod groups, with focus on mass extinctions and radiations: A phylogenetic approach".
- ^ a b c Tetlie 2007, p. 557.
- ^ Poschmann & Tetlie 2004, p. 189.
- ^ a b c d Braddy, Poschmann & Tetlie 2008, p. 107.
- ^ a b c Lamsdell & Braddy 2009, Supplementary information.
- ^ Briggs 1985, pp. 157–158.
- ^ Kjellesvig-Waering 1961, p. 830.
- ^ Lamsdell et al. 2015, p. 15.
- ^ Kraus & Brauckmann 2003, pp. 5–50.
- ^ Tetlie 2008, p. 19.
- ^ a b c d Tetlie 2007, p. 559.
- ^ a b c d Palaeos.
- ^ Whyte 2005, p. 576.
- ^ Selden 1999, p. 43.
- ^ Selden 1999, p. 45.
- ^ Selden 1999, pp. 44–46.
- ^ Hanken & Størmer 1975, pp. 262–267.
- ^ Braddy & Almond 1999, p. 166.
- ^ a b Braddy & Almond 1999, pp. 168–170.
- ^ Brezinski & Kollar 2016, p. 39.
- ^ Hanken & Størmer 1975, p. 255.
- ^ Vrazo & Ciurca 2017, p. 235.
- ^ Selden 1985, p. 219.
- ^ Selden 1985, pp. 220–221.
- ^ Selden 1985, p. 221.
- ^ Selden 1985, p. 222.
- ^ a b Selden 1985, pp. 222–223.
- ^ a b Lamsdell & Selden 2013, p. 32.
- ^ Lamsdell & Selden 2013, p. 33.
- ^ Lamsdell & Selden 2013, p. 44.
- ^ a b Lamsdell & Selden 2013, p. 34.
- ^ a b c Selden 1999, p. 46.
- ^ Selden 1999, p. 47.
- ^ Hembree, Platt & Smith 2014, p. 77.
- ^ Lamsdell, Braddy & Tetlie 2009, p. 1119.
- ^ Braddy & Dunlop 1997, p. 436.
- ^ Braddy & Dunlop 1997, p. 438.
- ^ Braddy, Poschmann & Tetlie 2008, p. 108.
- ^ Braddy & Dunlop 1997, p. 439.
- ^ Braddy & Dunlop 1997, p. 449.
- ^ Lamsdell 2014, pp. 175–177.
- ^ Braddy & Dunlop 1997, pp. 450–452.
- ^ Braddy & Dunlop 1997, pp. 454–455.
- ^ a b O'Connell 1916, p. 11.
- ^ a b Lamsdell et al. 2015, p. 1.
- ^ Van Roy, Briggs & Gaines 2015, p. 6.
- ^ a b c Lamsdell et al. 2015, p. 29.
- ^ O'Connell 1916, p. 12.
- ^ O'Connell 1916, p. 13.
- ^ Ortega-Hernández, Legg & Braddy 2012, p. 15.
- ^ Tetlie 2007, p. 569.
- ^ Tetlie 2007, p. 567.
- ^ a b c d e Tetlie 2007, p. 570.
- ^ Dunlop, Penney & Jekel 2018, pp. 17–30.
- ^ Selden 1999, p. 44.
- ^ Tetlie 2006, p. 410.
- ^ Tetlie & Rábano 2007, p. 124.
- ^ McCoy et al. 2015, p. 3.
- ^ Tetlie 2007, p. 571.
- ^ a b c Hallam & Wignall 1997, p. 70.
- ^ Lamsdell & Braddy 2009, p. 265.
- ^ a b Lamsdell & Braddy 2009, p. 266.
- ^ Lamsdell & Braddy 2009, p. 268.
- ^ Tetlie 2007.
- S2CID 59488956.
- S2CID 246819305.
- ^ Dunlop, Penney & Jekel 2018, pp. 19 & 24.
- ^ Tetlie & Van Roy 2006, p. 79.
- ^ Dunlop, Penney & Jekel 2018, p. 24.
- ^ Lamsdell et al. 2020, p. 1.
- ^ Kues & Kietzke 1981, p. 727.
- ^ Dunlop, Penney & Jekel 2018, p. 19.
- ^ White 1927, p. 575.
- ^ a b Poschmann & Rozefelds 2021.
- ^ Bergstrom & Dugatkin 2012, p. 515.
- ^ Clarke & Ruedemann 1912, p. 13.
- ^ Nudds & Selden 2008, pp. 78–82.
- ^ Burmeister 1843, pp. 62–64.
- ^ Dunlop, Penney & Jekel 2018, p. 27.
- ^ Kjellesvig-Waering 1964, p. 331.
- ^ M'Coy 1849, p. 393.
- ^ Henderson 1866, p. 18.
- ^ Clarke & Ruedemann 1912, p. 14.
- ^ Woodward 1865, pp. 484–486.
- ^ Clarke & Ruedemann 1912, p. 19.
- ^ Clarke & Ruedemann 1912, pp. 124–125.
- ^ Clarke & Ruedemann 1912, p. 135 & 137.
- ^ Clarke & Ruedemann 1912, p. 135.
- ^ a b Clarke & Ruedemann 1912, p. 136.
- ^ Clarke & Ruedemann 1912, p. 137.
- ^ Kingsley 1894, p. 119.
- ^ Lankester 1886, p. 366.
- ^ Dunlop, Penney & Jekel 2018, p. 17.
- ^ Lamsdell 2012, p. 19.
- ^ Lamsdell 2012, pp. 20–21.
- ^ Tollerton 1989, p. 642.
- ^ Tollerton 1989, pp. 642–644.
- ^ Tollerton 1989, p. 649.
- ^ Tollerton 1989, p. 646.
- ^ Tollerton 1989, p. 650.
- ^ Lamsdell et al. 2015, p. 25.
- ^ Tetlie & Cuggy 2007, p. 350.
- ^ Tetlie 2007, p. 565.
- ^ Lamsdell, Braddy & Tetlie 2010, p. 56.
- ^ Lamsdell et al. 2015, p. 3.
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Websites
- Kazlev, M. Alan (2002). "Palaeos - Eurypterida". www.palaeos.com. Archived from the original on 13 August 2007.
External links
- Eurypterids.co.uk – An online resource of eurypterid data and research
- eurypterid.net