1191:
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.
1492:
1473:
3739:
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). 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.
3735:
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. They are not applicable to all eurypterids either; stylonurine eurypterids lack swimming leg paddles entirely. 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. Most superfamilies and families are defined based on the morphology of the appendages.
3936:
1069:
4025:
3743:
members of the same suborder. 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. 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.
2387:
3700:
1142:
4232:
3689:
4205:
1634:
1587:
4323:
4549:
1247:
4667:
4156:
2571:
3868:
1703:, 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
2165:
1583:(used to deposit eggs). The different types of genital appendages are not necessarily the only feature that distinguishes between the sexes of eurypterids. Depending on the genus and species in question, other features such as size, the amount of ornamentation and the proportional width of the body can be the result of sexual dimorphism. In general, eurypterids with type B appendages (males) appear to have been proportionally wider than eurypterids with type A appendages (females) of the same genera.
4529:
177:
2035:, were the new apex predators in marine environments. 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. A recent review of Adelophthalmoidea admitted that "There is much more of a marine influence in many of the sections yielding
3978:
8453:
3800:
1092:(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
973:
3848:
2658:
Entomostraca, by later researchers such as John
Sterling Kinsgsley. In subsequent research, Gigantostraca has been treated as synonymous with Merostomata (rarely) and Eurypterida itself (more commonly). 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
1778:
4005:
142:
4252:
1915:
8756:
3916:
4469:
4653:
2002:
4502:
534:
4359:
945:
755:
2582:
1359:
1041:, 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
4111:
4379:
2441:, 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.
4433:
1275:(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
1206:. 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
1310:. 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.
999:, 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
910:, 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
1381:(the ability to perceive depth). The legs of many eurypterids were covered in thin spines, used both for locomotion and the gathering of food. In some groups, these spiny appendages became heavily specialized. In some eurypterids in the Carcinosomatoidea, forward-facing appendages were large and possessed enormously elongated spines (as in
1339:, 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.
2634:(the developmental stage immediately following the embryonic stage) in both groups, during which both xiphosurans and eurypterids have a proportionally larger carapace than adults, are generally broader, possess a distinct ridge down the middle, have a lesser number of segments which lack differentiation and have an underdeveloped telson.
1173:, 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
2657:
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
2641:
called
Merostomata (erected to house both groups by Henry Woodward in 1866). Though xiphosurans (like the eurypterids) were historically seen as crustaceans due to their respiratory system and their aquatic lifestyle, this hypothesis was discredited after numerous similarities were discovered between
1992:
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
825:
measured 36.4 centimeters (14.3 in) in length, but is missing a quarter of its length, suggesting that the full chelicera would have been 45.5 centimeters (17.9 in) long. If the proportions between body length and chelicerae match those of its closest relatives, where the ratio between claw
3742:
Modern research favors a classification into suborders
Eurypterina and Stylonurina instead, supported by phylogenetic analyses. In particular, pterygotid eurypterids share a number of homologies with derived eurypterine eurypterids such as the adelophthalmids, and are thus best classified as derived
3738:
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
2141:
is known from the terminal
Permian of Australia, which represents the youngest known eurypterid. 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.
1980:
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
1543:
where the type A appendage is divided into three but the type B appendage into only two. Such division of the genital appendage is common in eurypterids, but the number is not universal; for instance, the appendages of both types in the family
Pterygotidae are undivided. The type A appendage is also
1018:
due to a matching size (the trackmaker was estimated to have been about 1.6 meters (5.2 ft) long) and inferred leg anatomy. It is the largest terrestrial trackwayâmeasuring 6 meters (20 ft) long and averaging 95 centimeters (3.12 ft) in widthâmade by an arthropod found thus far. It is
3746:
The cladogram presented below, covering all currently recognized eurypterid families, follows a 2007 study by O. Erik Tetlie. The stylonurine suborder follows a 2010 study by James Lamsdell, Simon J. Braddy and Tetlie. The superfamily "Megalograptoidea", recognized by Tetlie in 2007 and then placed
2026:
Only three eurypterid familiesâAdelophthalmidae, Hibbertopteridae and Mycteroptidaeâsurvived 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., and with marine eurypterid predators
1809:
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. In most Silurian fossil beds, eurypterine eurypterids account for 90% of all eurypterids present.
1609:
on the type A appendages may have aided in breaking open the spermatophore to release the free sperm inside for uptake. The "horn organs," possibly spermathecae, are thought to have been connected directly to the appendage via tracts, but these supposed tracts remain unpreserved in available fossil
830:
that possessed the chelicera in question would have measured between 233 and 259 centimeters (7.64 and 8.50 ft), an average 2.5 meters (8.2 ft), in length. With the chelicerae extended, another meter (3.28 ft) would be added to this length. This estimate exceeds the maximum body size
3734:
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
2618:
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
1535:
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
1395:
members of the Pterygotioidea, the appendages were completely without spines, but had specialized claws instead. Other eurypterids, lacking these specialized appendages, likely fed in a manner similar to modern horseshoe crabs, by grabbing and shredding food with their appendages before pushing it
1889:
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
1696:
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. The earliest
1602:
rather than to serve as an ovipositor, as arthropod ovipositors are generally longer than eurypterid type A appendages. By rotating the sides of the operculum, it would have been possible to lower the appendage from the body. Due to the way different plates overlay at its location, the appendage
1227:
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
2087:
was widespread, living primarily in brackish and freshwater environments adjacent to coastal plains. These environments were maintained by favorable climate conditions. They did not persist as climate changes owing to Pangaea's formation altered depositional and vegetational patterns across the
1578:
received from males. This would imply that the type A appendage is the female morph and the type B appendage is the male. Further evidence for the type A appendages representing the female morph of genital appendages comes in their more complex construction (a general trend for female arthropod
1190:
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
1509:
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
1806:, 423 to 419.2 million years ago, of the very latest Silurian. This peak in diversity has been recognized since the early twentieth century; of the approximately 150 species of eurypterids known in 1916, more than half were from the Silurian and a third were from the Late Silurian alone.
1376:
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
1232:. 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
1027:
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
2494:
in which all eurypterid species thus far recovered from fossil deposits there were discussed. Clarke and Ruedemann created one of the first phylogenetic trees of eurypterids, dividing the order into two families; Eurypteridae (distinguished by smooth eyes and including
1613:
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
1536:
internal duct, is found in two distinct morphs, generally referred to as "type A" and "type B". These genital appendages are often preserved prominently in fossils and have been the subject of various interpretations of eurypterid reproduction and sexual dimorphism.
2629:
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. These ontogenetical similarities were seen as most apparent when studying the
1984:
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
1287:). 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.
2698:). Lamsdell noted that it is possible that Dekatriata is synonymous with Sclerophorata as the reproductive system, the primary defining feature of sclerophorates, has not been thoroughly studied in chasmataspidids. Dekatriata is, in turn, part of the
1881:
and Laurentia), which had been completely colonized by the genus during its merging and was unable to cross the vast expanses of ocean separating this continent from other parts of the world, such as the southern supercontinent Gondwana. As such,
2462:
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
740:
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.
705:
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
1019:
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
3751:
and Eurypteroidea, has been omitted as more recent studies suggest that the megalograptids were members of the superfamily Carcinosomatoidea. As such, the phylogeny of the Carcinosomatoidea follows a 2015 study by Lamsdell and colleagues.
618:(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
883:
measured 1.75 meters (5.7 ft), 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.
2454:(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. In the work
1346:, 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
1061:. 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
1023:, 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
729:
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
2060:
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. Out of the 33 species referred to
662:. 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 "
2043:
from the Carboniferous of New Mexico concluded that the habitat of some eurypterids "may need to be re-evaluated". The sole surviving eurypterine family, Adelophthalmidae, was represented by only a single genus,
831:
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;
1739:. Today only 11 species can be confidently identified as representing Ordovician eurypterids. These taxa fall into two distinct ecological categories; large and active predators from the ancient continent of
1971:
saw an additional five families going extinct. As marine groups were the most affected, the eurypterids were primarily impacted within the eurypterine suborder. Only one group of stylonurines (the family
919:
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
7265:
Poschmann, Markus; Rozefelds, Andrew (2021-11-30). "The last eurypterid â a southern high-latitude record of sweep-feeding sea scorpion from Australia constrains the timing of their extinction".
2686:
but not including all descendants of this ancestor) and thus not a valid phylogenetic group. Eurypterids were recovered as closely related to arachnids instead of xiphosurans, forming the group
1527:
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
1902:
itself, were active apex predators in Late Silurian marine ecosystems. The pterygotids were also evidently capable of crossing oceans, becoming one of only two eurypterid groups to achieve a
1224:(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.
1620:
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.
2625:) 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
673:
The underside of the opisthosoma was covered in structures evolved from modified opisthosomal appendages. Throughout the opisthosoma, these structures formed plate-like structures termed
598:
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
1710:
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
927:
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
4859:
7430:
6685:
Kues, Barry S.; Kietzke, Kenneth K. (1981). "A Large Assemblage of a New Eurypterid from the Red Tanks Member, Madera Formation (Late Pennsylvanian-Early Permian) of New Mexico".
1121:
preserves the largest eurypterid footprints known to date with the found tracks each being about 7.6 centimeters (3.0 in) in diameter. Other eurypterid ichnogenera include
2273:, within a clade he named "Palaeadae". Within Palaeadae, Burmeister erected three families; the "Trilobitae" (composed of all trilobites), the "Cytherinidae" (composed only of
1735:
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
1322:
491:, 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,
1810:
Though some were likely already present by the Late Ordovician (simply missing from the fossil record so far), a vast majority of eurypterid groups are first recorded in
1566:
associated with the type A appendages is a set of organs traditionally described as either "tubular organs" or "horn organs". These organs are most often interpreted as
1459:
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.
1103:
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
7989:
1714:
age, they are not recognized as eurypterids, and sometimes not even as related forms, today. Some animals previously seen as primitive eurypterids, such as the genus
1099:
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
1655:
Until 1882 no eurypterids were known from before the Silurian. Contemporary discoveries since the 1880s have expanded the knowledge of early eurypterids from the
887:
Giant eurypterids were not limited to the family Pterygotidae. An isolated 12.7 centimeters (5.0 in) long fossil metastoma of the carcinosomatoid eurypterid
1111:, the discoveries of trackways both predate and outnumber eurypterid body fossils. Eurypterid trackways have been referred to several ichnogenera, most notably
625:, 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.
2344:" by the workers. Agassiz first thought the fossils represented remains of fish, only recognizing their nature as arthropod remains five years later in 1844.
7181:
Ortega-HernĂĄndez, Javier; Legg, David A.; Braddy, Simon J. (2012). "The phylogeny of aglaspidid arthropods and the internal relationships within Artiopoda".
497:, 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
2263:. He considered the trilobites to be crustaceans, as previous authors had, and classified them together with what he assumed to be their closest relatives,
2553:. In line with earlier authors, Clarke and Ruedemann also supported a close relationship between the eurypterids and the horseshoe crabs (united under the
2467:
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
1963:
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
6670:"Fossil giants and surviving dwarfs. Arthropleurida and Pselaphognatha (Atelocerata, Diplopoda): characters, phylogenetic relationships and construction"
670:" (comprising segments 7 to 12) or into a "preabdomen" (generally comprising segments 1 to 7) and "postabdomen" (generally comprising segments 8 to 12).
2365:(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
1603:
would have been impossible to move without muscular contractions moving around the operculum. It would have been kept in place when not it use. The
606:
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
2646:
in 1894, classified the Merostomata as a sister group to the Arachnida under the class "Acerata" within a subphylum "Branchiata". Others, such as
8860:
1944:
Though the eurypterids continued to be abundant and diversify during the Early Devonian (for instance leading to the evolution of the pterygotid
1203:
1146:
7640:, a Stylonurid (Chelicerata: Eurypterida) from the Late Devonian Catskill Delta Complex, and Its Phylogenetic Position in the Hardieopteridae".
1981:
affected during the Late Devonian, when many of the older groups were replaced by new forms in the families Mycteroptidae and Hibbertopteridae.
7598:
1510:
morphology alone. Sometimes two sexes of the same species have been interpreted as two different species, as was the case with two species of
7376:
3708:
The most important taxonomic character in eurypterids is the morphology of the sixth pair of appendages. In most eurypterids of the suborder
2127:, which might have reached 1.4 meters (4.6 ft) in length, while originally considered to be Late Permian in age other sources suggest a
1802:
Eurypterids were most diverse and abundant between the Middle Silurian and the Early Devonian, with an absolute peak in diversity during the
1539:
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
4874:
3279:
893:
indicates the animal would have reached a length of 2.2 meters (7.2 ft) in life, rivalling the pterygotids in size. Another giant was
7481:
1993:
within the eurypterids, which gave rise to several new forms capable of "sweep-feeding" (raking through the substrate in search of prey).
7302:
Poschmann, Markus; Tetlie, O. Erik (2004). "On the Emsian (Early Devonian) arthropods of the Rhenish Slate Mountains: 4. The eurypterids
2269:
1003:. In contrast, a number of stylonurines had elongated and powerful legs that might have allowed them to walk on land (similar to modern
3294:
1472:
4805:"Selectivity in the evolution of Palaeozoic arthropod groups, with focus on mass extinctions and radiations: A phylogenetic approach"
2475:). The description also helped solidify the close relationship between the eurypterids and other chelicerates by showcasing numerous
931:
would possibly have rivalled the largest pterygotids in weight, if not surpassed them, and as such be among the heaviest arthropods.
156:
1598:
The primary function of the long, assumed female, type A appendages was likely to take up spermatophore from the substrate into the
1491:
1427:
was opened through the thin cuticle between the last segment before the telson and the telson itself, as in modern horseshoe crabs.
7102:"All the better to see you with: eyes and claws reveal the evolution of divergent ecological roles in giant pterygotid eurypterids"
6987:
Selectivity in the evolution of Palaeozoic arthropod groups, with focus on mass extinctions and radiations: a phylogenetic approach
1769:), are likely to represent the first truly successful eurypterid group, experiencing a small radiation during the Late Ordovician.
1212:. Instead, among arthropod respiratory organs, the eurypterid gill tracts most closely resemble the pseudotracheae found in modern
632:, the posteriormost division of the body, which in most species took the form of a blade-like shape. In some lineages, notably the
7560:
Tetlie, O. Erik; Cuggy, Michael B. (2007). "Phylogeny of the basal swimming eurypterids (Chelicerata; Eurypterida; Eurypterina)".
7077:"XLI.âOn the classification of some British fossil Crustacea, with notices of new forms in the University Collection at Cambridge"
2123:
persisted longer. A massive incomplete carapace from Permian deposits in Russia represents the sole fossil remains of the species
1117:(defined as a series of four tracks often with an associated drag mark in the mid-line), wherein the holotype of the ichnospecies
1556:
in the type B appendage is also possible and the structure may represent the unfused tips of the appendages. Located between the
6997:"The oldest described eurypterid: a giant Middle Ordovician (Darriwilian) megalograptid from the Winneshiek Lagerstätte of Iowa"
8929:
8532:
8522:
8044:
7411:
7154:
6302:
Braddy, Simon J.; Almond, John E. (1999). "Eurypterid trackways from the Table Mountain Group (Ordovician) of South Africa".
2143:
2099:
Mycteroptids and hibbertopterids continued to survive for some time, with one genus of each group known from Permian strata:
2069:
had already been relatively widespread and represented around all major landmasses in the Late Devonian, the amalgamation of
416:
407:
accounts for more than 90% of all known eurypterid specimens. Though the group continued to diversify during the subsequent
7997:
5711:"A new species of the eurypterid Hibbertopterus from the Carboniferous of New Mexico, and a review of the Hibbertopteridae"
1890:
appeared. Though the largest members of the family appeared in the Devonian, large two meter (6.5+ ft) pterygotids such as
1857:) account for more than 90% (perhaps as many as 95%) of all known fossil eurypterid specimens. Despite their vast number,
1430:
Eurypterid coprolites discovered in deposits of Ordovician age in Ohio containing fragments of a trilobite and eurypterid
1014:
was discovered in Carboniferous-aged fossil deposits of Scotland in 2005. It was attributed to the stylonurine eurypterid
397:
order. Following their appearance during the Ordovician, eurypterids became major components of marine faunas during the
8484:
6542:
6483:
6446:
6253:
2255:
published his view on trilobite taxonomy and how the group related to other organisms, living and extinct, in the work
6647:
Kjellesvig-Waering, Erik N. (1964). "A Synopsis of the Family Pterygotidae Clarke and Ruedemann, 1912 (Eurypterida)".
2650:
in 1909, went further and classified the Merostomata as a subclass within the Arachnida, raised to the rank of class.
8914:
8499:
6521:
6408:
2019:
1838:
2244:
2065:, 23 (69%) are from the Carboniferous alone. The genus reached its peak diversity in the Late Carboniferous. Though
1732:. The aglaspidids, once seen as primitive chelicerates, are now seen as a group more closely related to trilobites.
470:
7714:
Tollerton, Victor P. (1989). "Morphology, Taxonomy, and Classification of the Order Eurypterida Burmeister, 1843".
8017:
2212:
examined the fossil and recognized it as clearly belonging to an arthropod. He thought the fossil, which he named
8743:
8527:
8469:
2220:, and suggested it might represent a missing link between the trilobites and more derived branchiopods. The name
2159:
1167:
In eurypterids, the respiratory organs were located on the ventral body wall (the underside of the opisthosoma).
644:, the telson was flattened and may have been used as a rudder while swimming. Some genera within the superfamily
7669:
Tetlie, O. Erik; Briggs, Derek E. G. (2009). "The origin of pterygotid eurypterids (Chelicerata: Eurypterida)".
2232:
1976:) went extinct in the Early Devonian. Only two families of eurypterines survived into the Late Devonian at all (
614:(claws). The subsequent pairs of appendages, numbers II to VI, possessed gnathobases (or "tooth-plates") on the
456:
8934:
8517:
8507:
8461:
6708:"Cope's Rule and Romer's theory: patterns of diversity and gigantism in eurypterids and Palaeozoic vertebrates"
1434:
in association with full specimens of the same eurypterid species have been suggested to represent evidence of
412:
1960:-building organisms) effectively crippled the abundance and diversity previously seen within the eurypterids.
8479:
8474:
7653:
1194:
It has been suggested instead that the "gill tract" was an organ for breathing air, perhaps actually being a
2135:
age (~283-267 million years ago) for the specimen. A fragment of eurypterid cuticle, given the species name
1574:), though this function is yet to be proven conclusively. In arthropods, spermathecae are used to store the
799:. Sizes around 100 centimeters (3.3 ft) are common in most eurypterid groups. The smallest eurypterid,
8512:
8489:
8011:
1894:
were already present during the Late Silurian. Their ecology ranged from generalized predatory behavior to
482:), meaning 'wing', referring to the pair of wide swimming appendages present in many members of the group.
7100:
McCoy, Victoria E.; Lamsdell, James C.; Poschmann, Markus; Anderson, Ross P.; Briggs, Derek E. G. (2015).
6532:
2637:
Due to these similarities, the xiphosurans and eurypterids have often been united under a single class or
8924:
2438:
1177:(small spines), which resulted in an enlarged surface area. It was composed of spongy tissue due to many
795:
Eurypterids were highly variable in size, depending on factors such as lifestyle, living environment and
572:
509:. Only a handful of eurypterid groups spread beyond the confines of Euramerica and a few genera, such as
17:
1755:
and Gondwana. The Laurentian predators, classified in the family Megalograptidae (comprising the genera
1579:
genitalia). It is possible that the greater length of the type A appendage means that it was used as an
176:
2445:
899:, a primitive carcinosomatoid, which is estimated to have reached lengths of 1.7 meters (5.6 ft).
7760:
7603:(Chelicerata, Eurypterida) in the collections of Museo Geominero (Geological Survey of Spain), Madrid"
446:
was present, which would have allowed for short periods of time in terrestrial environments. The name
8145:
8037:
4680:
1903:
1392:
1318:
1267:
Like all arthropods, eurypterids matured and grew through static developmental stages referred to as
522:
6374:
Brezinski, David K.; Kollar, Albert D. (2016). "Reevaluation of the Age and Provenance of the Giant
7625:
7001:
6935:
1949:
1929:
7399:
6995:
Lamsdell, James C.; Briggs, Derek E. G.; Liu, Huaibao; Witzke, Brian J.; McKay, Robert M. (2015).
6473:
4024:
8939:
7761:"The Fezouata fossils of Morocco; an extraordinary record of marine life in the Early Ordovician"
7340:
3935:
2595:
2472:
2195:
902:
Typical of large eurypterids is a lightweight build. Factors such as locomotion, energy costs in
810:
2598:(top), united in the class Merostomata. Recent studies are in favor of a closer relationship to
2150:
recorded, and rendered many other successful Paleozoic groups, such as the trilobites, extinct.
1342:
The most frequently observed change occurring through ontogeny (except for some genera, such as
505:
because the group lived primarily in the waters around and within the ancient supercontinent of
8891:
8774:
8452:
7951:
Woodward, Henry (1865). "On some New Species of Crustacea belonging to the Order Eurypterida".
7228:
Plotnick, Roy E.; Baumiller, Tomasz K. (1988). "The pterygotid telson as a biological rudder".
6436:
2643:
2557:
Merostomata) but also discussed alternative hypotheses such as a closer relation to arachnids.
2380:
2209:
2191:
2175:
1068:
723:
552:
2073:
into a global supercontinent over the course of the last two periods of the Paleozoic allowed
8886:
8878:
8812:
7144:
5656:"Phylogeny and palaeoecology of the Adelophthalmoidea (Arthropoda; Chelicerata; Eurypterida)"
2511:
712:
or "gill tracts", would potentially have aided eurypterids to breathe air above water, while
8865:
6904:"Revised systematics of Palaeozoic 'horseshoe crabs' and the myth of monophyletic Xiphosura"
6849:
Lamsdell, James C.; McCoy, Victoria E.; Perron-Feller, Opal A.; Hopkins, Melanie J. (2020).
485:
The eurypterid order includes the largest known arthropods ever to have lived. The largest,
8919:
8834:
8759:
8030:
7913:
7772:
7723:
7678:
7569:
7532:
7523:
Tetlie, O. Erik (2007). "Distribution and dispersal history of Eurypterida (Chelicerata)".
7449:
7355:
7237:
7183:
7010:
6944:
6864:
6811:
6762:
6494:
6457:
6311:
5722:
5667:
4672:
2683:
2386:
2301:
1199:
2433:, and an exhaustive description of the various eurypterids of New York in Volume 3 of the
1127:(though it is likely that many specimens actually represent trackways of crustaceans) and
8:
6800:"The systematics and phylogeny of the Stylonurina (Arthropoda: Chelicerata: Eurypterida)"
3699:
2430:
2426:
1378:
1150:
1037:
The functionality of the eurypterine swimming paddles varied from group to group. In the
796:
7917:
7776:
7727:
7682:
7573:
7536:
7453:
7359:
7241:
7014:
6948:
6868:
6815:
6766:
6626:
Kjellesvig-Waering, Erik N. (1961). "The Silurian Eurypterida of the Welsh Borderland".
6315:
5726:
5671:
401:, from which the majority of eurypterid species have been described. The Silurian genus
7968:
7939:
7881:
7854:
7841:
7806:
7747:
7739:
7702:
7657:
7585:
7473:
7327:
7290:
7249:
7216:
7146:
Fossil Ecosystems of North America: A Guide to the Sites and their Extraordinary Biotas
7126:
7101:
7033:
6996:
6967:
6930:
6890:
6837:
6786:
6732:
6707:
6694:
6669:
6656:
6635:
6614:
6606:
6577:
6395:
6357:
6332:
6285:
6264:
5746:
5691:
4204:
3748:
2638:
2476:
2252:
2187:
2170:
2164:
1861:
are only known from a relatively short temporal range, first appearing during the Late
1675:
1599:
1046:
956:
in which the shape of the paddles and their motion through water is enough to generate
610:, the chelicerae were large and long, with strong, well-developed teeth on specialised
603:
443:
320:
249:
171:
151:
7964:
7500:
6323:
4231:
3688:
1290:
Hemianamorphic direct development has been observed in many arthropod groups, such as
1141:
8873:
8348:
8281:
8273:
7972:
7931:
7886:
7845:
7810:
7798:
7694:
7690:
7617:
7548:
7465:
7417:
7391:
7331:
7294:
7282:
7253:
7208:
7200:
7196:
7150:
7131:
7038:
6972:
6920:
6903:
6894:
6882:
6829:
6790:
6778:
6774:
6737:
6581:
6538:
6517:
6479:
6442:
6362:
6290:
6249:
5750:
5738:
5683:
4658:
4322:
4295:
4151:
2483:
2394:
2390:
1633:
1417:. Though a potential anal opening has been reported from the telson of a specimen of
1093:
645:
7751:
7706:
7661:
7589:
7477:
7220:
7051:
6841:
6618:
6399:
5695:
4694:âfamily of fishes in which some members contain "sea scorpion" in their common name.
4548:
1948:, the largest of all arthropods), the group was one of many heavily affected by the
1886:
was limited geographically to the coastlines and shallow inland seas of Euramerica.
8688:
8402:
8394:
8384:
8376:
8356:
8327:
8243:
8192:
7960:
7943:
7921:
7904:
7876:
7866:
7831:
7788:
7780:
7731:
7686:
7649:
7577:
7540:
7457:
7363:
7319:
7274:
7245:
7192:
7121:
7113:
7088:
7063:
7028:
7018:
6962:
6952:
6915:
6872:
6819:
6770:
6753:(Chelicerata: Eurypterida: Stylonurina) from the late Devonian of Portishead, UK".
6727:
6719:
6598:
6569:
6509:
6387:
6352:
6344:
6319:
6280:
5730:
5675:
4464:
4428:
4354:
4247:
4000:
3843:
3718:(top), this leg is modified into a swimming paddle. In eurypterids of the suborder
2487:
2398:
2183:
2147:
2006:
1977:
1973:
1952:. The extinction event, only known to affect marine life (particularly trilobites,
1862:
1827:
1557:
1400:
1246:
1123:
1062:
637:
383:
7278:
6985:
6534:
Experimental Approaches to Understanding Fossil Organisms: Lessons from the Living
5734:
1865:(around 432 million years ago) and being extinct by the end of the Pridoli epoch.
1145:
The supposed "gill tracts" of eurypterids have been compared to the air-breathing
30:
This article is about the order Eurypterida. For the eurypterid family containing
8717:
8563:
8420:
8361:
8317:
8220:
8207:
7544:
7435:(Chelicerata: Eurypterida) from Norway and Canada and the phylogeny of the genus"
6855:
6425:
6243:
4697:
4497:
4318:
4200:
4155:
3973:
3911:
3897:
3867:
3511:
2786:
2695:
2674:
2554:
2348:
2182:
The first known eurypterid specimen was discovered in the Silurian-aged rocks of
2137:
2107:
2010:
1895:
1819:
1660:
1648:
1586:
1347:
1271:. These instars were punctuated by periods during which eurypterids went through
1011:
818:
364:
6749:
Lamsdell, James C.; Braddy, Simon J.; Tetlie, O. Erik (2009). "Redescription of
6513:
5710:
2300:
in 1838, not identified as eurypterids until later), out of those still seen as
1329:, composed of multiple specimens of various developmental stages of eurypterids
1306:. True direct development has on occasion been referred to as a trait unique to
340:
8633:
8591:
8549:
8412:
8339:
8299:
8291:
8215:
7859:
Proceedings of the National Academy of Sciences of the United States of America
6851:"Air Breathing in an Exceptionally Preserved 340-Million-Year-Old Sea Scorpion"
4528:
4482:
4283:
4106:
3931:
3724:
2631:
2523:
2501:
2459:
2046:
1920:
1480:
1335:
1259:
1030:
980:
843:
808:
The largest eurypterid, and the largest known arthropod ever to have lived, is
719:
633:
615:
511:
431:
7735:
7581:
7461:
7092:
7067:
7023:
6877:
6850:
6824:
6799:
5793:
5679:
426:
Although popularly called "sea scorpions", only the earliest eurypterids were
8908:
8640:
8612:
8605:
8577:
8556:
8309:
8248:
8238:
8230:
8184:
8174:
8166:
8122:
8086:
7802:
7698:
7621:
7552:
7469:
7395:
7286:
7257:
7204:
6833:
6782:
6294:
5742:
5687:
4185:
4020:
3977:
3958:
3829:
3795:
3524:
3412:
3385:
3199:
3028:
2875:
2687:
2654:
2647:
2609:
2604:
2517:
2325:
2039:
than has previously been acknowledged." Similarly, a study of the eurypterid
2028:
1933:
1849:. Thought to have hunted mainly small and soft-bodied invertebrates, such as
1823:
1815:
1803:
1757:
1744:
1699:
1639:
1575:
1512:
1456:
1440:
1419:
1387:
1220:(windpipes) of air-breathing organisms, are lung-like and present within the
1038:
895:
788:
766:
695:
in German). These created a branchial chamber (gill tract) between preceding
641:
611:
588:
498:
487:
451:
387:
331:
309:
231:
92:
7505:
Stainier, 1917 and its position within the Adelophthalmidae Tollerton, 1989"
7165:
6957:
5655:
3799:
1826:
as well as eurypterine groups such as the Pterygotioidea, Eurypteroidea and
390:
period. With approximately 250 species, the Eurypterida is the most diverse
8797:
8598:
8430:
8366:
8322:
8197:
7935:
7890:
7793:
7367:
7212:
7135:
7117:
7042:
6976:
6886:
6741:
6723:
6573:
6366:
6348:
5555:
4524:
4374:
4227:
3863:
3847:
3367:
3224:
2954:
2679:
2619:
appendages bearing the "gill tracts" on appendages of the opisthosoma (the
2570:
2545:
2366:
2282:
2217:
2032:
1796:
1783:
1736:
1499:
1445:
1229:
1178:
1104:
1050:
1042:
957:
912:
833:
801:
607:
568:
493:
415:. They declined in numbers and diversity until becoming extinct during the
130:
54:
35:
7871:
4004:
1777:
1216:. These organs, called pseudotracheae, because of some resemblance to the
972:
8847:
8806:
8695:
8672:
8570:
8264:
8157:
8095:
8077:
6391:
4424:
4251:
4083:
3772:
3719:
3714:
3709:
3482:
3238:
3161:
3075:
3058:
2937:
2925:
2893:
2769:
2752:
2703:
2699:
2668:
2549:). Both families were considered to be descended from a common ancestor,
1925:
1725:
1711:
1679:
1667:
1567:
1435:
1113:
1078:
996:
985:
961:
907:
889:
782:
619:
580:
435:
427:
394:
371:
301:
280:
276:
266:
218:
141:
67:
50:
8826:
7784:
6589:
Kingsley, John Sterling (1894). "The Classification of the Arthropoda".
6427:
Die Organisation der Trilobiten aus ihren lebenden Verwandten entwickelt
6000:
5757:
5709:
Braddy, Simon J.; Lerner, Allan J; Lucas, Spencer G. (1 February 2023).
5450:
5416:
5414:
5412:
4468:
3915:
2198:
in 1818. He erroneously identified the fossil as an example of the fish
1914:
916:, and are possibly vital for the evolution of giant size in arthropods.
8661:
8647:
8626:
8619:
8584:
8425:
8113:
8104:
7836:
7819:
7743:
7421:
7413:
Treatise on Invertebrate Paleontology, Part P Arthropoda 2, Chelicerata
7323:
6698:
6660:
6639:
6610:
5387:
5385:
4544:
4501:
3496:
3455:
3343:
3255:
3108:
2978:
2833:
2731:
2714:
2691:
2471:
was comparable with the knowledge of its modern relatives (such as the
2450:
2093:
2081:
2001:
1986:
1953:
1870:
1846:
1729:
1656:
1580:
1413:
1363:
1299:
1295:
1284:
1088:
1054:
952:
874:
864:
862:, the Pterygotidae, is noted for several unusually large species. Both
772:
760:
599:
539:
517:
506:
403:
375:
112:
77:
31:
8852:
7348:
Philosophical Transactions of the Royal Society B: Biological Sciences
6171:
4849:
4847:
4845:
4843:
4804:
4652:
4358:
1853:, species of the genus (of which the most common is the type species,
1833:
The most successful eurypterid by far was the Middle to Late Silurian
1313:
Studies on a well-preserved fossil assemblage of eurypterids from the
1049:) in which the motion and shape of the paddles are enough to generate
8738:
8722:
8712:
8071:
6219:
5409:
4904:
4688:âan unrelated family of insects, commonly known as "water scorpions".
3538:
3182:
3126:
3044:
2996:
2599:
2591:
2361:
2258:
2237:
2225:
2128:
2101:
1968:
1842:
1740:
1687:
1686:, but these have yet to be thoroughly studied, and are likely to be
1303:
1291:
1280:
1000:
650:
548:
533:
476:
462:
391:
360:
208:
188:
160:
117:
61:
8768:
7926:
7899:
7076:
6553:
5382:
5295:
5259:
1358:
1045:. Larger individuals may have been capable of underwater flying (or
944:
8821:
8791:
8681:
8654:
6602:
6207:
4840:
4691:
2803:
2581:
2371:
2337:
2278:
1964:
1814:
of Silurian age. These include both stylonurine groups such as the
1792:
1788:
1752:
1748:
1721:
1716:
1704:
1674:. There are also reports of even earlier fossil eurypterids in the
1403:
have been reported from fossils of various eurypterids, among them
1369:
1307:
1276:
1221:
1129:
1108:
1073:
1058:
754:
731:
667:
663:
655:
622:
584:
439:
408:
398:
107:
102:
87:
82:
72:
5988:
5507:
5055:
5053:
4110:
2375:, based on fossil remains previously assigned to a new species of
906:
and respiration, as well as the actual physical properties of the
6848:
6265:"The functional morphology of mating in the Silurian eurypterid,
6029:
6027:
5916:
5846:
5844:
5799:
4892:
4775:
4773:
4745:
4743:
4741:
4739:
4737:
4735:
4733:
4731:
4729:
4685:
3437:
2205:
2200:
2132:
2070:
1937:
1878:
1874:
1811:
1683:
1326:
1314:
1272:
1217:
1213:
1174:
1159:
1154:
903:
855:
size estimate is based on trackway evidence, not fossil remains.
822:
734:
margin of this structure, the genital appendage (also called the
702:
576:
560:
556:
164:
122:
97:
8839:
7512:
Bulletin de l'Institut Royal des Sciences Naturelles de Belgique
6931:"Babes in the wood â a unique window into sea scorpion ontogeny"
5856:
5613:
5611:
5397:
5026:
4378:
2706:(the only monophyletic xiphosuran group) and other stem-genera.
2531:) and Pterygotidae (distinguished by faceted eyes and including
1096:
might have been adjustable by raising and positioning the tail.
8065:
7759:
Van Roy, Peter; Briggs, Derek E. G.; Gaines, Robert R. (2015).
6075:
5904:
5817:
5781:
5355:
5343:
5247:
5208:
5206:
5169:
5167:
5050:
4916:
4432:
2850:
2341:
1268:
1254:
826:
size and body length is relatively consistent, the specimen of
815:
629:
592:
564:
502:
382:. The group is likely to have appeared first either during the
198:
8022:
7099:
6798:
Lamsdell, James C.; Braddy, Simon J.; Tetlie, O. Erik (2010).
6039:
6024:
6012:
5841:
5586:
5584:
5582:
5561:
5372:
5370:
4770:
4760:
4758:
4726:
2590:
Eurypterids have historically been seen as closely related to
2208:-like appearance of the carapace. Seven years later, in 1825,
2186:, to this day one of the richest eurypterid fossil locations.
7820:"New trace fossil evidence for eurypterid swimming behaviour"
6331:
Braddy, Simon J.; Poschmann, Markus; Tetlie, O. Erik (2008).
6123:
5976:
5952:
5868:
5623:
5608:
5596:
5065:
4866:
4864:
4862:
4828:
4785:
2678:) concluded that the Xiphosura, as presently understood, was
2642:
the horseshoe crabs and the arachnids. Some authors, such as
1571:
1531:
and the eighth segment (distinctly plate-like) is called the
1250:
659:
6674:
Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg
5880:
5203:
5191:
5179:
5164:
5154:
5152:
5077:
6531:
Hembree, Daniel I.; Platt, Brian F.; Smith, Jon J. (2014).
6183:
5964:
5892:
5769:
5647:
5579:
5543:
5438:
5426:
5367:
5331:
5319:
5307:
5283:
5271:
5038:
4755:
2336:
and when the first fossils were discovered by quarrymen in
1957:
1850:
1424:
1195:
1133:(which preserves grooves made by the swimming appendages).
1004:
805:, measured just 2.03 centimeters (0.80 in) in length.
7180:
7052:"Professor Claus and the classification of the Arthropoda"
6099:
5805:
5456:
5089:
1065:, using this type of propulsion is more energy-efficient.
6159:
6147:
6135:
6111:
5149:
5113:
4880:
1671:
379:
7654:
10.3374/0079-032X(2008)49[19:HEASCE]2.0.CO;2
6063:
5928:
5497:
5495:
5493:
5491:
5489:
5225:
5223:
5221:
4949:
4947:
4945:
4943:
7170:
The Bulletin of the Buffalo Society of Natural Sciences
6994:
6456:
Dunlop, Jason A.; Penney, David; Jekel, Denise (2018).
6225:
6177:
6051:
5940:
5420:
5391:
5014:
4910:
4818:
4816:
4814:
2292:
The fourth eurypterid genus to be described (following
2092:
dwindled in number and had already gone extinct by the
587:(sometimes called the "prosomal shield") on which both
6458:"A summary list of fossil spiders and their relatives"
6195:
6087:
2369:, closely related to horseshoe crabs. A fourth genus,
6378:
Eurypterid Trackway, from Elk County, Pennsylvania".
6330:
5654:
Tetlie, O. Erik; Poschmann, Markus (1 January 2008).
5567:
5531:
5486:
5474:
5462:
5301:
5218:
5137:
5125:
5101:
4940:
4853:
1643:, the earliest known eurypterid. The family to which
411:
period, the eurypterids were heavily affected by the
8014:â An online resource of eurypterid data and research
6797:
6748:
6213:
5829:
5519:
5265:
5235:
5002:
4990:
4978:
4928:
4811:
4714:
4648:
1082:, containing the largest eurypterid footprints known
7758:
5403:
758:Size comparison of six of the largest eurypterids:
555:bodies and jointed appendages (limbs) covered in a
7855:"Flora of the Hermit Shale, Grand Canyon, Arizona"
7264:
6646:
6625:
5922:
5850:
5702:
5635:
4968:
4966:
4964:
4962:
4898:
1928:(with swimming paddles) eurypterid to survive the
1659:period. The earliest eurypterids known today, the
1651:, was the first truly successful eurypterid group.
7818:Vrazo, Matthew B.; Ciurca, Samuel J. Jr. (2017).
7642:Bulletin of the Peabody Museum of Natural History
7525:Palaeogeography, Palaeoclimatology, Palaeoecology
7227:
6530:
6455:
6241:
6081:
5910:
5862:
5823:
5787:
5763:
5708:
5513:
5253:
4779:
1989:(armored fish) in both North America and Europe.
1444:from the Ordovician of Ohio contain fragments of
8906:
6562:Transactions of the Edinburgh Geological Society
2413:(now seen as synonymous with another species of
1967:stage four families went extinct, and the later
658:and may have been capable of using it to inject
7301:
6667:
6434:
6373:
6333:"Giant claw reveals the largest ever arthropod"
6242:Bergstrom, Carl T.; Dugatkin, Lee Alan (2012).
6045:
6033:
6018:
6006:
5994:
5982:
5958:
5874:
5653:
5071:
4959:
4922:
4834:
4791:
4147:
4102:
3791:
1228:forced water into the invaginations leading to
595:(simple eye-like sensory organs) were located.
6928:
6705:
6435:Clarke, John Mason; Ruedemann, Rudolf (1912).
5629:
5617:
5602:
5212:
5197:
5185:
5173:
4870:
2379:, was referred to the Eurypteridae in 1856 by
995:The two eurypterid suborders, Eurypterina and
628:The opisthosoma comprised 12 segments and the
8038:
7498:
7163:
6706:Lamsdell, James C.; Braddy, Simon J. (2009).
6492:
6471:
5775:
5590:
5444:
5432:
5376:
5083:
5032:
1438:. Similar coprolites referred to the species
370:. The earliest known eurypterids date to the
7668:
7596:
6301:
6262:
5549:
5361:
5349:
5325:
5313:
5289:
5277:
5059:
5044:
4764:
4749:
2620:
2409:(1858) featured an extensive description of
1996:
1561:
1233:
1207:
1185:
1168:
735:
713:
707:
696:
674:
7953:Quarterly Journal of the Geological Society
7817:
7559:
7142:
6684:
6493:Hanken, Nils-Martin; Størmer, Leif (1975).
6263:Braddy, Simon J.; Dunlop, Jason A. (1997).
6189:
5886:
5811:
5095:
2309:
1615:
1604:
1551:
1545:
680:
8045:
8031:
6558:, from the Silurian of the Pentland Hills"
6495:"The trail of a large Silurian eurypterid"
6472:Hallam, Anthony; Wignall, Paul B. (1997).
6423:
5898:
2115:went extinct during the Early Permian, as
2077:to gain an almost worldwide distribution.
1107:and the rest of the former supercontinent
923:measures 65 cm (26 in) wide. As
140:
7925:
7880:
7870:
7835:
7792:
7713:
7125:
7074:
7049:
7032:
7022:
6966:
6956:
6929:Lamsdell, James C.; Selden, Paul (2013).
6919:
6908:Zoological Journal of the Linnean Society
6876:
6823:
6731:
6668:Kraus, Otto; Brauckmann, Carsten (2003).
6551:
6356:
6284:
6273:Zoological Journal of the Linnean Society
6165:
6153:
6141:
6129:
6117:
6069:
5946:
5934:
3678:
1396:into their mouth using their chelicerae.
960:. This type of locomotion was limited to
654:, had a telson similar to that of modern
157:State Museum of Natural History Karlsruhe
7950:
7597:Tetlie, O. Erik; RĂĄbano, Isabel (2007).
7499:Tetlie, O. Erik; Van Roy, Peter (2006).
6983:
6901:
6588:
6105:
6093:
6057:
5970:
5457:Ortega-HernĂĄndez, Legg & Braddy 2012
5337:
2456:Anatomy and Relations of the Eurypterida
2385:
2261:aus ihren lebenden Verwandten entwickelt
2216:, represented a crustacean of the order
2190:described the specimen, discovered near
2163:
2050:. The hibbertopterids, mycteroptids and
2000:
1913:
1776:
1751:) and basal animals from the continents
1632:
1585:
1550:(lit. 'fork' in Latin). The presence of
1357:
1245:
1140:
1067:
1034:, was probably faster and more precise.
753:
583:(abdomen). The prosoma was covered by a
532:
7409:
4720:
2144:Permian–Triassic extinction event
2014:, the last known surviving eurypterid.
1623:
1462:
872:measured 2.1 meters (6.9 ft), and
722:, would cover the parts that serve for
14:
8907:
7900:"A gigantic fossil arthropod trackway"
7635:
7522:
7428:
7374:
7338:
7081:Annals and Magazine of Natural History
7056:Annals and Magazine of Natural History
6441:. University of California Libraries.
6406:
6201:
5641:
5573:
5537:
5525:
5501:
5480:
5468:
5241:
5229:
5158:
5143:
5131:
5119:
5107:
5020:
5008:
4996:
4953:
4934:
4886:
4822:
1670:stage of the Middle Ordovician, 467.3
1448:and fragments of smaller specimens of
8773:
8772:
8026:
7897:
7852:
7410:Størmer, Leif (1955). "Merostomata".
7143:Nudds, John R.; Selden, Paul (2008).
5835:
4984:
2332:was considerably larger in size than
1869:was also restricted to the continent
988:(lacking swimming paddles) eurypterid
950:Illustration of subaqueous flight in
7377:"Autecology of Silurian Eurypterids"
7310:n. gen. (Arthropoda: Chelicerata)".
6475:Mass Extinctions and Their Aftermath
6409:"Gigantism in Palaeozoic arthropods"
1841:, equally likely to have engaged in
1787:, a member of the highly successful
1544:armed with two curved spines called
964:(with swimming paddles) eurypterids.
666:" (comprising segments 1 to 6) and "
525:with fossils being found worldwide.
7987:
7562:Journal of Systematic Palaeontology
7442:Journal of Systematic Palaeontology
6804:Journal of Systematic Palaeontology
6537:. Springer Science & Business.
5660:Journal of Systematic Palaeontology
5302:Braddy, Poschmann & Tetlie 2008
4972:
4854:Braddy, Poschmann & Tetlie 2008
2421:), which, along with the monograph
2153:
442:. Some studies suggest that a dual
24:
7250:10.1111/j.1502-3931.1988.tb01746.x
6286:10.1111/j.1096-3642.1997.tb01282.x
6267:Baltoeurypterus tetragonophthalmus
6214:Lamsdell, Braddy & Tetlie 2010
5266:Lamsdell, Braddy & Tetlie 2009
2080:During the Late Carboniferous and
468:), meaning 'broad' or 'wide', and
419:(or sometime shortly before) 251.9
25:
8951:
8005:
7965:10.1144/GSL.JGS.1865.021.01-02.52
7765:Journal of the Geological Society
6304:Journal of African Earth Sciences
5404:Van Roy, Briggs & Gaines 2015
2608:(bottom), united under the clade
2560:
2289:, then including three species).
2020:Permian-Triassic extinction event
1238:, remain unknown in eurypterids.
417:PermianâTriassic extinction event
8755:
8754:
8451:
7691:10.1111/j.1475-4983.2009.00907.x
7197:10.1111/j.1096-0031.2012.00413.x
7166:"The Habitat of the Eurypterida"
6921:10.1111/j.1096-3642.2012.00874.x
6775:10.1111/j.1475-4983.2009.00902.x
4665:
4651:
4547:
4527:
4500:
4467:
4431:
4377:
4357:
4321:
4250:
4230:
4203:
4154:
4109:
4023:
4003:
3976:
3934:
3914:
3866:
3846:
3798:
3698:
3687:
2694:(composed of sclerophorates and
2580:
2569:
2088:world. With their habitat gone,
1932:and persist into the subsequent
1490:
1471:
971:
943:
571:, the body was divided into two
175:
65:
8744:Timeline of eurypterid research
8052:
7384:Special Papers in Palaeontology
6413:Special Papers in Palaeontology
6234:
6082:Dunlop, Penney & Jekel 2018
5911:Dunlop, Penney & Jekel 2018
5824:Dunlop, Penney & Jekel 2018
5788:Dunlop, Penney & Jekel 2018
5764:Dunlop, Penney & Jekel 2018
5514:Dunlop, Penney & Jekel 2018
5254:Hembree, Platt & Smith 2014
2160:Timeline of eurypterid research
7416:. University of Kansas Press.
6467:. Natural History Museum Bern.
5851:Poschmann & Rozefelds 2021
4797:
2393:of eurypterids as imagined by
1697:known stylonurine eurypterid,
1136:
718:, similar to organs in modern
528:
413:Late Devonian extinction event
13:
1:
8930:Darriwilian first appearances
7431:"Two new Silurian species of
7279:10.1080/08912963.2021.1998033
6324:10.1016/S0899-5362(99)00087-1
5863:Bergstrom & Dugatkin 2012
5735:10.1080/08912963.2022.2032690
4780:Plotnick & Baumiller 1988
4703:
2277:, an animal today seen as an
2204:, likely due to the strange,
1423:, it is more likely that the
1053:, similar to the swimming of
934:
844:Hibbertopterus wittebergensis
7545:10.1016/j.palaeo.2007.05.011
7164:O'Connell, Marjorie (1916).
6424:Burmeister, Hermann (1843).
6407:Briggs, Derek E. G. (1985).
4708:
2238:
2226:
2096:stage of the Early Permian.
1590:Type A genital appendage of
1560:and ventral surfaces of the
1497:Type B genital appendage of
1478:Type A genital appendage of
1441:Lanarkopterus dolichoschelus
1253:(left) and juvenile (right)
851:is very fragmentary and the
430:; many later forms lived in
27:Order of arthropods (fossil)
7:
7981:
6984:Lamsdell, James C. (2014).
6902:Lamsdell, James C. (2012).
6514:10.18261/8200049639-1975-16
6478:. Oxford University Press.
6438:The Eurypterida of New York
6046:Clarke & Ruedemann 1912
6034:Clarke & Ruedemann 1912
6019:Clarke & Ruedemann 1912
6007:Clarke & Ruedemann 1912
5995:Clarke & Ruedemann 1912
5983:Clarke & Ruedemann 1912
5959:Clarke & Ruedemann 1912
5875:Clarke & Ruedemann 1912
5072:Brezinski & Kollar 2016
4923:Kraus & Brauckmann 2003
4835:Poschmann & Tetlie 2004
4792:Clarke & Ruedemann 1912
4644:
2492:The Eurypterida of New York
2142:This extinction event, the
2138:Woodwardopterus freemanorum
2054:survived into the Permian.
2022:or sometime shortly before.
1909:
1772:
1241:
1086:Some eurypterines, such as
10:
8956:
7610:BoletĂn GeolĂłgico y Minero
7050:Lankester, E. Ray (1886).
5630:Lamsdell & Braddy 2009
5618:Lamsdell & Braddy 2009
5603:Lamsdell & Braddy 2009
5213:Lamsdell & Selden 2013
5198:Lamsdell & Selden 2013
5186:Lamsdell & Selden 2013
5174:Lamsdell & Selden 2013
4871:Lamsdell & Braddy 2009
2340:they were referred to as "
2243:
2231:
2157:
2146:, is the most devastating
1682:(Early Ordovician) age in
1628:
1500:Kokomopterus longicaudatus
1353:
1279:in some lineages, such as
896:Pentecopterus decorahensis
802:Alkenopterus burglahrensis
767:Pentecopterus decorahensis
744:
469:
455:
355:, often informally called
29:
8781:
8752:
8731:
8705:
8671:
8541:
8498:
8460:
8449:
8411:
8393:
8375:
8347:
8336:
8308:
8290:
8272:
8261:
8229:
8206:
8183:
8165:
8154:
8146:List of eurypterid genera
8138:
8131:
8060:
7898:Whyte, Martin A. (2005).
7736:10.1017/S0022336000041275
7636:Tetlie, O. Erik (2008). "
7582:10.1017/S1477201907002131
7462:10.1017/S1477201906001921
7093:10.1080/03745486009494858
7075:M'Coy, Frederick (1849).
7068:10.1080/00222938609460154
7024:10.1186/s12862-015-0443-9
6878:10.1016/j.cub.2020.08.034
6825:10.1080/14772011003603564
6380:Annals of Carnegie Museum
5776:Tetlie & Van Roy 2006
5680:10.1017/S1477201907002416
5591:Hallam & Wignall 1997
5084:Hanken & Størmer 1975
5033:Hanken & Størmer 1975
4875:Supplementary information
4681:List of eurypterid genera
4542:
4522:
4515:
4495:
4487:
4462:
4455:
4426:
4418:
4410:
4372:
4352:
4345:
4316:
4308:
4300:
4288:
4245:
4225:
4218:
4198:
4190:
4178:
4149:
4141:
4133:
4104:
4096:
4088:
4018:
3998:
3991:
3971:
3963:
3929:
3909:
3902:
3890:
3861:
3841:
3834:
3822:
3793:
3785:
3777:
3765:
3551:
3536:
3529:
3508:
3501:
3478:
3451:
3433:
3426:
3408:
3381:
3363:
3356:
3348:
3292:
3276:
3269:
3251:
3243:
3220:
3213:
3195:
3187:
3175:
3157:
3149:
3122:
3104:
3097:
3089:
3071:
3063:
3040:
3033:
2992:
2975:
2968:
2950:
2942:
2930:
2889:
2871:
2864:
2846:
2838:
2826:
2799:
2783:
2765:
2757:
2745:
2727:
2719:
2621:
2435:Palaeontology of New York
1997:Carboniferous and Permian
1904:cosmopolitan distribution
1616:
1605:
1562:
1552:
1481:Adelophthalmus mansfieldi
1319:Beartooth Butte Formation
1234:
1208:
1169:
773:Acutiramus macrophthalmus
761:Pterygotus grandidentatus
714:
697:
523:cosmopolitan distribution
438:, and they were not true
359:, are a group of extinct
326:
319:
262:
257:
172:Scientific classification
170:
148:
139:
45:
8915:Prehistoric chelicerates
7988:Kazlev, M. Alan (2002).
7429:Tetlie, O. Erik (2006).
7341:"Eurypterid respiration"
7002:BMC Evolutionary Biology
6936:BMC Evolutionary Biology
6552:Henderson, John (1866).
6376:Palmichnium kosinskiorum
6009:, p. 135 & 137.
5550:Tetlie & RĂĄbano 2007
5362:Braddy & Dunlop 1997
5350:Braddy & Dunlop 1997
5326:Braddy & Dunlop 1997
5314:Braddy & Dunlop 1997
5290:Braddy & Dunlop 1997
5278:Braddy & Dunlop 1997
5060:Braddy & Almond 1999
5045:Braddy & Almond 1999
4765:Tetlie & Briggs 2009
4750:Braddy & Dunlop 1997
2702:, a group including the
2479:between the two groups.
2018:went extinct during the
1950:Late Devonian extinction
1930:Late Devonian extinction
1724:, are now classified as
575:(sections); the frontal
551:, eurypterids possessed
543:with body parts labelled
7990:"Palaeos - Eurypterida"
7716:Journal of Paleontology
7312:Senckenbergiana Lethaea
6990:. University of Kansas.
6958:10.1186/1471-2148-13-98
6751:Drepanopterus abonensis
6687:Journal of Paleontology
6649:Journal of Paleontology
6628:Journal of Paleontology
6591:The American Naturalist
6190:Tetlie & Cuggy 2007
5923:Kjellesvig-Waering 1964
5887:Nudds & Selden 2008
5812:Kues & Kietzke 1981
5766:, pp. 19 & 24.
5096:Vrazo & Ciurca 2017
4899:Kjellesvig-Waering 1961
2596:Atlantic horseshoe crab
2473:Atlantic horseshoe crab
2423:On the Genus Pterygotus
1432:Megalograptus ohioensis
1079:Palmichnium kosinkiorum
868:, whose largest member
814:. A chelicera from the
811:Jaekelopterus rhenaniae
789:Jaekelopterus rhenaniae
749:
7368:10.1098/rstb.1985.0081
7118:10.1098/rsbl.2015.0564
6724:10.1098/rsbl.2009.0700
6574:10.1144/transed.1.1.15
6349:10.1098/rsbl.2007.0491
3679:Internal relationships
2644:John Sterling Kingsley
2402:
2179:
2023:
2005:Reconstruction of the
1941:
1799:
1652:
1595:
1546:
1373:
1264:
1186:
1164:
1083:
921:Hibbertoperus scouleri
879:whose largest species
792:
736:
724:underwater respiration
708:
675:
544:
8935:Lopingian extinctions
8887:Paleobiology Database
7872:10.1073/pnas.13.8.574
7853:White, David (1927).
7638:Hallipterus excelsior
7375:Selden, Paul (1999).
7339:Selden, Paul (1985).
7149:. Manson Publishing.
2419:E. tetragonophthalmus
2407:De Euryptero Remipede
2389:
2304:in modern times, was
2257:Die Organisation der
2167:
2004:
1917:
1877:continents Avalonia,
1780:
1720:from the Cambrian of
1636:
1589:
1361:
1249:
1144:
1071:
890:Carcinosoma punctatum
783:Carcinosoma punctatum
757:
579:(head) and posterior
536:
343:& Waterston, 1968
6465:World Spider Catalog
6392:10.2992/007.084.0105
6226:Lamsdell et al. 2015
6178:Lamsdell et al. 2015
5800:Lamsdell et al. 2020
5421:Lamsdell et al. 2015
5392:Lamsdell et al. 2015
4911:Lamsdell et al. 2015
4673:Palaeontology portal
3728:(bottom), it is not.
2684:last common ancestor
2448:described the genus
1637:A reconstruction of
1624:Evolutionary history
1570:(organs for storing
1463:Reproductive biology
837:(closely related to
834:Erettopterus grandis
7918:2005Natur.438..576W
7785:10.1144/jgs2015-017
7777:2015JGSoc.172..541V
7728:1989JPal...63..642T
7683:2009Palgy..52.1141T
7574:2007JSPal...5..345T
7537:2007PPP...252..557T
7454:2006JSPal...4..397T
7360:1985RSPTB.309..219S
7242:1988Letha..21...13P
7015:2015BMCEE..15..169L
6949:2013BMCEE..13...98L
6869:2020CBio...30E4316L
6816:2010JSPal...8...49L
6767:2009Palgy..52.1113L
6316:1999JAfES..29..165B
6132:, pp. 642â644.
5997:, pp. 124â125.
5973:, pp. 484â486.
5727:2023HBio...35..257B
5672:2008JSPal...6..237T
5364:, pp. 454â455.
5352:, pp. 450â452.
5340:, pp. 175â177.
5161:, pp. 222â223.
5122:, pp. 220â221.
5062:, pp. 168â170.
5035:, pp. 262â267.
4889:, pp. 157â158.
4752:, pp. 437â439.
2682:(a group sharing a
2465:Eurypterus fischeri
2431:John William Salter
2427:Thomas Henry Huxley
2411:Eurypterus fischeri
2302:taxonomically valid
2236:'broad, wide') and
2224:derives from Greek
1399:Fossils preserving
1379:stereoscopic vision
423:million years ago.
149:Fossil specimen of
8925:Chelicerate orders
8274:Onychopterelloidea
8000:on 13 August 2007.
7837:10.1111/pala.12336
7503:Eurypterus dumonti
7501:"A reappraisal of
7405:on August 3, 2011.
7324:10.1007/BF03043470
7267:Historical Biology
6556:Slimonia Acuminata
6502:Fossils and Strata
5715:Historical Biology
3749:Onychopterelloidea
2405:Jan Nieszkowski's
2403:
2253:Hermann Burmeister
2214:Eurypterus remipes
2188:Samuel L. Mitchill
2180:
2171:Eurypterus remipes
2024:
1942:
1918:Reconstruction of
1898:and some, such as
1800:
1795:eurypterid family
1781:Reconstruction of
1653:
1600:reproductive tract
1596:
1374:
1265:
1181:in the structure.
1165:
1084:
853:H. wittenbergensis
797:taxonomic affinity
793:
545:
444:respiratory system
152:Eurypterus remipes
8902:
8901:
8874:Open Tree of Life
8775:Taxon identifiers
8766:
8765:
8447:
8446:
8443:
8442:
8439:
8438:
8395:Adelophthalmoidea
8377:Waeringopteroidea
8349:Carcinosomatoidea
8282:Onychopterellidae
8257:
8256:
8012:Eurypterids.co.uk
7631:on July 22, 2011.
7354:(1138): 219â226.
7273:(10): 2020â2030.
7156:978-1-84076-088-0
6863:(21): 4316â4321.
6108:, pp. 20â21.
5901:, pp. 62â64.
5889:, pp. 78â82.
5562:McCoy et al. 2015
5516:, pp. 17â30.
5023:, pp. 44â46.
4659:Arthropods portal
4641:
4640:
4632:
4631:
4623:
4622:
4614:
4613:
4605:
4604:
4596:
4595:
4587:
4586:
4578:
4577:
4569:
4568:
4560:
4559:
4444:
4443:
4399:
4398:
4390:
4389:
4334:
4333:
4296:Carcinosomatoidea
4272:
4271:
4263:
4262:
4167:
4166:
4152:Onychopterellidae
4122:
4121:
4072:
4071:
4063:
4062:
4054:
4053:
4045:
4044:
4036:
4035:
3947:
3946:
3879:
3878:
3811:
3810:
3675:
3674:
3664:
3663:
3655:
3654:
3646:
3645:
3637:
3636:
3628:
3627:
3619:
3618:
3610:
3609:
3601:
3600:
3592:
3591:
3583:
3582:
3574:
3573:
3565:
3564:
3467:
3466:
3397:
3396:
3332:
3331:
3323:
3322:
3314:
3313:
3305:
3304:
3138:
3137:
3017:
3016:
3008:
3007:
2914:
2913:
2905:
2904:
2815:
2814:
2690:within the clade
2484:John Mason Clarke
2395:John Mason Clarke
2391:Evolutionary tree
1873:(composed of the
1828:Waeringopteroidea
1672:million years ago
1367:depicted hunting
1323:Cottonwood Canyon
1094:center of gravity
1047:subaqueous flight
881:P. grandidentatus
646:Carcinosomatoidea
380:million years ago
350:
349:
344:
335:
294:
284:
271:
253:
134:
60:467.3â251.9
16:(Redirected from
8947:
8895:
8894:
8882:
8881:
8869:
8868:
8856:
8855:
8843:
8842:
8830:
8829:
8817:
8816:
8815:
8802:
8801:
8800:
8770:
8769:
8758:
8757:
8732:Related articles
8689:Merostomichnites
8455:
8403:Adelophthalmidae
8385:Waeringopteridae
8357:Carcinosomatidae
8345:
8344:
8328:Strobilopteridae
8270:
8269:
8244:Hibbertopteridae
8193:Parastylonuridae
8163:
8162:
8136:
8135:
8047:
8040:
8033:
8024:
8023:
8001:
7996:. Archived from
7976:
7959:(1â2): 484â486.
7947:
7929:
7894:
7884:
7874:
7849:
7839:
7814:
7796:
7755:
7710:
7677:(5): 1141â1148.
7665:
7632:
7630:
7624:. Archived from
7607:
7593:
7556:
7531:(3â4): 557â574.
7519:
7509:
7495:
7493:
7492:
7486:
7480:. Archived from
7439:
7425:
7406:
7404:
7398:. Archived from
7381:
7371:
7345:
7335:
7318:(1â2): 173â193.
7298:
7261:
7224:
7177:
7160:
7139:
7129:
7096:
7071:
7062:(100): 364â372.
7046:
7036:
7026:
6991:
6980:
6970:
6960:
6925:
6923:
6898:
6880:
6845:
6827:
6794:
6761:(5): 1113â1139.
6745:
6735:
6702:
6681:
6664:
6643:
6622:
6597:(326): 118â135.
6585:
6548:
6527:
6499:
6489:
6468:
6462:
6452:
6431:
6420:
6403:
6370:
6360:
6327:
6298:
6288:
6269:(Fischer, 1839)"
6259:
6229:
6223:
6217:
6211:
6205:
6199:
6193:
6187:
6181:
6175:
6169:
6163:
6157:
6151:
6145:
6139:
6133:
6127:
6121:
6115:
6109:
6103:
6097:
6091:
6085:
6079:
6073:
6067:
6061:
6055:
6049:
6043:
6037:
6031:
6022:
6016:
6010:
6004:
5998:
5992:
5986:
5980:
5974:
5968:
5962:
5956:
5950:
5944:
5938:
5932:
5926:
5920:
5914:
5908:
5902:
5896:
5890:
5884:
5878:
5872:
5866:
5860:
5854:
5848:
5839:
5833:
5827:
5821:
5815:
5809:
5803:
5797:
5791:
5785:
5779:
5773:
5767:
5761:
5755:
5754:
5706:
5700:
5699:
5651:
5645:
5639:
5633:
5627:
5621:
5615:
5606:
5600:
5594:
5588:
5577:
5571:
5565:
5559:
5553:
5547:
5541:
5535:
5529:
5523:
5517:
5511:
5505:
5499:
5484:
5478:
5472:
5466:
5460:
5454:
5448:
5442:
5436:
5430:
5424:
5418:
5407:
5401:
5395:
5389:
5380:
5374:
5365:
5359:
5353:
5347:
5341:
5335:
5329:
5323:
5317:
5311:
5305:
5299:
5293:
5287:
5281:
5275:
5269:
5263:
5257:
5251:
5245:
5239:
5233:
5227:
5216:
5210:
5201:
5195:
5189:
5183:
5177:
5171:
5162:
5156:
5147:
5141:
5135:
5129:
5123:
5117:
5111:
5105:
5099:
5093:
5087:
5081:
5075:
5069:
5063:
5057:
5048:
5042:
5036:
5030:
5024:
5018:
5012:
5006:
5000:
4994:
4988:
4982:
4976:
4970:
4957:
4951:
4938:
4932:
4926:
4925:, pp. 5â50.
4920:
4914:
4908:
4902:
4896:
4890:
4884:
4878:
4868:
4857:
4851:
4838:
4832:
4826:
4820:
4809:
4808:
4801:
4795:
4789:
4783:
4777:
4768:
4762:
4753:
4747:
4724:
4718:
4675:
4670:
4669:
4668:
4661:
4656:
4655:
4551:
4531:
4518:
4517:
4504:
4490:
4489:
4471:
4465:Adelophthalmidae
4458:
4457:
4435:
4429:Waeringopteridae
4421:
4420:
4413:
4412:
4381:
4361:
4355:Carcinosomatidae
4348:
4347:
4325:
4311:
4310:
4303:
4302:
4291:
4290:
4254:
4248:Strobilopteridae
4234:
4221:
4220:
4207:
4193:
4192:
4181:
4180:
4158:
4144:
4143:
4136:
4135:
4113:
4099:
4098:
4091:
4090:
4027:
4007:
4001:Hibbertopteridae
3994:
3993:
3980:
3966:
3965:
3938:
3918:
3905:
3904:
3893:
3892:
3870:
3850:
3844:Parastylonuridae
3837:
3836:
3825:
3824:
3802:
3788:
3787:
3780:
3779:
3768:
3767:
3756:
3755:
3702:
3691:
3532:
3531:
3504:
3503:
3429:
3428:
3359:
3358:
3351:
3350:
3272:
3271:
3246:
3245:
3216:
3215:
3190:
3189:
3178:
3177:
3152:
3151:
3100:
3099:
3092:
3091:
3066:
3065:
3036:
3035:
2971:
2970:
2945:
2944:
2933:
2932:
2867:
2866:
2841:
2840:
2829:
2828:
2760:
2759:
2748:
2747:
2722:
2721:
2710:
2709:
2624:
2623:
2584:
2573:
2488:Rudolf Ruedemann
2399:Rudolf Ruedemann
2324:), described by
2323:
2320:
2317:
2314:
2311:
2247:
2241:
2235:
2229:
2154:History of study
1978:Adelophthalmidae
1974:Parastylonuridae
1896:ambush predation
1863:Llandovery epoch
1666:, date from the
1619:
1618:
1608:
1607:
1565:
1564:
1555:
1554:
1549:
1494:
1475:
1401:digestive tracts
1237:
1236:
1211:
1210:
1189:
1172:
1171:
1124:Merostomichnites
1063:drag coefficient
978:Illustration of
975:
947:
739:
717:
716:
711:
700:
699:
694:
691:
688:
685:
682:
678:
638:Hibbertopteridae
480:
473:
466:
459:
422:
384:Early Ordovician
339:
330:
290:
279:
270:Burmeister, 1843
269:
248:
243:
180:
179:
144:
128:
127:
64:
49:Temporal range:
43:
42:
21:
8955:
8954:
8950:
8949:
8948:
8946:
8945:
8944:
8905:
8904:
8903:
8898:
8890:
8885:
8877:
8872:
8864:
8859:
8851:
8846:
8838:
8833:
8825:
8820:
8811:
8810:
8805:
8796:
8795:
8790:
8777:
8767:
8762:
8748:
8727:
8718:Chasmataspidida
8701:
8667:
8564:Campylocephalus
8537:
8494:
8456:
8435:
8421:Hughmilleriidae
8407:
8389:
8371:
8362:Megalograptidae
8338:
8332:
8318:Dolichopteridae
8304:
8292:Moselopteroidea
8286:
8263:
8253:
8239:Drepanopteridae
8225:
8221:Hardieopteridae
8208:Kokomopteroidea
8202:
8179:
8156:
8150:
8127:
8056:
8051:
8008:
7994:www.palaeos.com
7984:
7979:
7927:10.1038/438576a
7794:1854/LU-8714212
7628:
7605:
7507:
7490:
7488:
7484:
7437:
7402:
7379:
7343:
7157:
7112:(8): 20150564.
7106:Biology Letters
7087:(24): 392â414.
6856:Current Biology
6712:Biology Letters
6554:"IV. Notice of
6545:
6524:
6497:
6486:
6460:
6449:
6430:. Georg Reimer.
6337:Biology Letters
6256:
6237:
6232:
6224:
6220:
6212:
6208:
6200:
6196:
6188:
6184:
6176:
6172:
6164:
6160:
6152:
6148:
6140:
6136:
6128:
6124:
6116:
6112:
6104:
6100:
6092:
6088:
6080:
6076:
6068:
6064:
6056:
6052:
6044:
6040:
6032:
6025:
6017:
6013:
6005:
6001:
5993:
5989:
5981:
5977:
5969:
5965:
5957:
5953:
5945:
5941:
5933:
5929:
5921:
5917:
5909:
5905:
5899:Burmeister 1843
5897:
5893:
5885:
5881:
5873:
5869:
5861:
5857:
5849:
5842:
5834:
5830:
5822:
5818:
5810:
5806:
5798:
5794:
5786:
5782:
5774:
5770:
5762:
5758:
5707:
5703:
5652:
5648:
5640:
5636:
5628:
5624:
5616:
5609:
5601:
5597:
5589:
5580:
5572:
5568:
5560:
5556:
5548:
5544:
5536:
5532:
5524:
5520:
5512:
5508:
5500:
5487:
5479:
5475:
5467:
5463:
5455:
5451:
5443:
5439:
5431:
5427:
5419:
5410:
5402:
5398:
5390:
5383:
5375:
5368:
5360:
5356:
5348:
5344:
5336:
5332:
5324:
5320:
5312:
5308:
5300:
5296:
5288:
5284:
5276:
5272:
5268:, p. 1119.
5264:
5260:
5252:
5248:
5240:
5236:
5228:
5219:
5211:
5204:
5196:
5192:
5184:
5180:
5172:
5165:
5157:
5150:
5142:
5138:
5130:
5126:
5118:
5114:
5106:
5102:
5094:
5090:
5082:
5078:
5070:
5066:
5058:
5051:
5043:
5039:
5031:
5027:
5019:
5015:
5007:
5003:
4995:
4991:
4983:
4979:
4971:
4960:
4952:
4941:
4933:
4929:
4921:
4917:
4909:
4905:
4897:
4893:
4885:
4881:
4869:
4860:
4852:
4841:
4833:
4829:
4821:
4812:
4803:
4802:
4798:
4790:
4786:
4778:
4771:
4767:, p. 1141.
4763:
4756:
4748:
4727:
4719:
4715:
4711:
4706:
4698:History of life
4671:
4666:
4664:
4657:
4650:
4647:
4642:
4633:
4624:
4615:
4606:
4597:
4588:
4579:
4570:
4561:
4498:Hughmilleriidae
4445:
4400:
4391:
4335:
4319:Megalograptidae
4273:
4264:
4201:Dolichopteridae
4168:
4123:
4073:
4064:
4055:
4046:
4037:
3974:Drepanopteridae
3948:
3912:Hardieopteridae
3898:Kokomopteroidea
3880:
3812:
3732:
3731:
3730:
3729:
3705:
3704:
3703:
3694:
3693:
3692:
3681:
3676:
3670:
3665:
3656:
3647:
3638:
3629:
3620:
3611:
3602:
3593:
3584:
3575:
3566:
3512:Chasmataspidida
3468:
3398:
3333:
3324:
3315:
3306:
3139:
3018:
3009:
2915:
2906:
2816:
2787:Trilobitomorpha
2696:chasmataspidids
2675:Stoermeropterus
2632:nepionic stages
2616:
2615:
2614:
2613:
2587:
2586:
2585:
2576:
2575:
2574:
2563:
2349:Frederick M'Coy
2321:
2318:
2315:
2312:
2298:Campylocephalus
2176:James E. De Kay
2162:
2156:
2148:mass extinction
2121:Campylocephalus
2108:Campylocephalus
2029:sarcopterygians
2016:Campylocephalus
2011:Campylocephalus
1999:
1912:
1820:Kokomopteroidea
1775:
1747:(living on the
1649:Megalograptidae
1631:
1626:
1507:
1506:
1505:
1504:
1503:
1495:
1486:
1485:
1484:
1476:
1465:
1356:
1348:Moselopteroidea
1244:
1153:legs of modern
1149:present in the
1139:
1012:fossil trackway
993:
992:
991:
990:
989:
976:
967:
966:
965:
948:
937:
821:of Willwerath,
819:Klerf Formation
752:
747:
720:horseshoe crabs
692:
689:
686:
683:
547:Like all other
537:Restoration of
531:
450:comes from the
420:
315:
296:
295:
272:
247:
241:
174:
135:
126:
125:
120:
115:
110:
105:
100:
95:
90:
85:
80:
75:
70:
59:
58:
47:
39:
28:
23:
22:
15:
12:
11:
5:
8953:
8943:
8942:
8940:Apex predators
8937:
8932:
8927:
8922:
8917:
8900:
8899:
8897:
8896:
8883:
8870:
8857:
8844:
8831:
8818:
8803:
8787:
8785:
8779:
8778:
8764:
8763:
8753:
8750:
8749:
8747:
8746:
8741:
8735:
8733:
8729:
8728:
8726:
8725:
8720:
8715:
8709:
8707:
8706:Related groups
8703:
8702:
8700:
8699:
8692:
8685:
8677:
8675:
8669:
8668:
8666:
8665:
8658:
8651:
8644:
8637:
8634:Onychopterella
8630:
8623:
8616:
8609:
8602:
8595:
8592:Hibbertopterus
8588:
8581:
8574:
8567:
8560:
8553:
8550:Adelophthalmus
8545:
8543:
8542:Notable genera
8539:
8538:
8536:
8535:
8530:
8525:
8520:
8515:
8510:
8504:
8502:
8496:
8495:
8493:
8492:
8487:
8482:
8477:
8472:
8466:
8464:
8458:
8457:
8450:
8448:
8445:
8444:
8441:
8440:
8437:
8436:
8434:
8433:
8428:
8423:
8417:
8415:
8413:Pterygotioidea
8409:
8408:
8406:
8405:
8399:
8397:
8391:
8390:
8388:
8387:
8381:
8379:
8373:
8372:
8370:
8369:
8364:
8359:
8353:
8351:
8342:
8340:Diploperculata
8334:
8333:
8331:
8330:
8325:
8320:
8314:
8312:
8306:
8305:
8303:
8302:
8300:Moselopteridae
8296:
8294:
8288:
8287:
8285:
8284:
8278:
8276:
8267:
8259:
8258:
8255:
8254:
8252:
8251:
8246:
8241:
8235:
8233:
8227:
8226:
8224:
8223:
8218:
8216:Kokomopteridae
8212:
8210:
8204:
8203:
8201:
8200:
8195:
8189:
8187:
8181:
8180:
8178:
8177:
8171:
8169:
8167:Rhenopteroidea
8160:
8152:
8151:
8149:
8148:
8142:
8140:
8133:
8129:
8128:
8126:
8125:
8116:
8107:
8098:
8089:
8080:
8074:
8068:
8061:
8058:
8057:
8050:
8049:
8042:
8035:
8027:
8021:
8020:
8018:eurypterid.net
8015:
8007:
8006:External links
8004:
8003:
8002:
7983:
7980:
7978:
7977:
7948:
7895:
7865:(8): 574â575.
7850:
7830:(2): 235â252.
7815:
7771:(5): 541â549.
7756:
7722:(5): 642â657.
7711:
7666:
7633:
7616:(1): 117â126.
7599:"Specimens of
7594:
7568:(3): 345â356.
7557:
7520:
7496:
7448:(4): 397â412.
7426:
7407:
7372:
7336:
7299:
7262:
7225:
7178:
7161:
7155:
7140:
7097:
7072:
7047:
6992:
6981:
6926:
6899:
6846:
6795:
6746:
6718:(2): 265â269.
6703:
6693:(4): 709â729.
6682:
6665:
6655:(2): 331â361.
6644:
6634:(4): 789â835.
6623:
6603:10.1086/275878
6586:
6549:
6544:978-9401787208
6543:
6528:
6522:
6490:
6485:978-0198549161
6484:
6469:
6453:
6448:978-1125460221
6447:
6432:
6421:
6404:
6371:
6343:(1): 106â109.
6328:
6310:(1): 165â177.
6299:
6279:(4): 435â461.
6260:
6255:978-0393913415
6254:
6238:
6236:
6233:
6231:
6230:
6218:
6206:
6204:, p. 565.
6194:
6192:, p. 350.
6182:
6170:
6168:, p. 650.
6166:Tollerton 1989
6158:
6156:, p. 646.
6154:Tollerton 1989
6146:
6144:, p. 649.
6142:Tollerton 1989
6134:
6130:Tollerton 1989
6122:
6120:, p. 642.
6118:Tollerton 1989
6110:
6098:
6086:
6074:
6072:, p. 366.
6070:Lankester 1886
6062:
6060:, p. 119.
6050:
6048:, p. 137.
6038:
6036:, p. 136.
6023:
6021:, p. 135.
6011:
5999:
5987:
5975:
5963:
5951:
5947:Henderson 1866
5939:
5937:, p. 393.
5927:
5925:, p. 331.
5915:
5903:
5891:
5879:
5867:
5865:, p. 515.
5855:
5840:
5838:, p. 575.
5828:
5816:
5814:, p. 727.
5804:
5792:
5780:
5768:
5756:
5721:(2): 257â263.
5701:
5666:(2): 237â249.
5646:
5634:
5632:, p. 268.
5622:
5620:, p. 266.
5607:
5605:, p. 265.
5595:
5578:
5576:, p. 571.
5566:
5554:
5552:, p. 124.
5542:
5540:, p. 410.
5530:
5518:
5506:
5504:, p. 570.
5485:
5483:, p. 567.
5473:
5471:, p. 569.
5461:
5449:
5445:O'Connell 1916
5437:
5433:O'Connell 1916
5425:
5408:
5396:
5381:
5377:O'Connell 1916
5366:
5354:
5342:
5330:
5328:, p. 449.
5318:
5316:, p. 439.
5306:
5304:, p. 108.
5294:
5292:, p. 438.
5282:
5280:, p. 436.
5270:
5258:
5246:
5234:
5217:
5202:
5190:
5178:
5163:
5148:
5146:, p. 222.
5136:
5134:, p. 221.
5124:
5112:
5110:, p. 219.
5100:
5098:, p. 235.
5088:
5086:, p. 255.
5076:
5064:
5049:
5047:, p. 166.
5037:
5025:
5013:
5001:
4989:
4987:, p. 576.
4977:
4958:
4956:, p. 559.
4939:
4927:
4915:
4903:
4901:, p. 830.
4891:
4879:
4858:
4856:, p. 107.
4839:
4837:, p. 189.
4827:
4825:, p. 557.
4810:
4796:
4794:, p. 244.
4784:
4769:
4754:
4725:
4712:
4710:
4707:
4705:
4702:
4701:
4700:
4695:
4689:
4683:
4677:
4676:
4662:
4646:
4643:
4639:
4638:
4635:
4634:
4630:
4629:
4626:
4625:
4621:
4620:
4617:
4616:
4612:
4611:
4608:
4607:
4603:
4602:
4599:
4598:
4594:
4593:
4590:
4589:
4585:
4584:
4581:
4580:
4576:
4575:
4572:
4571:
4567:
4566:
4563:
4562:
4558:
4557:
4554:
4553:
4541:
4538:
4537:
4534:
4533:
4521:
4516:
4514:
4511:
4510:
4507:
4506:
4494:
4488:
4486:
4485:
4483:Pterygotioidea
4478:
4477:
4474:
4473:
4461:
4456:
4454:
4451:
4450:
4447:
4446:
4442:
4441:
4438:
4437:
4425:
4419:
4417:
4411:
4409:
4406:
4405:
4402:
4401:
4397:
4396:
4393:
4392:
4388:
4387:
4384:
4383:
4371:
4368:
4367:
4364:
4363:
4351:
4346:
4344:
4341:
4340:
4337:
4336:
4332:
4331:
4328:
4327:
4315:
4309:
4307:
4301:
4299:
4298:
4289:
4287:
4286:
4284:Diploperculata
4279:
4278:
4275:
4274:
4270:
4269:
4266:
4265:
4261:
4260:
4257:
4256:
4244:
4241:
4240:
4237:
4236:
4224:
4219:
4217:
4214:
4213:
4210:
4209:
4197:
4191:
4189:
4188:
4179:
4177:
4174:
4173:
4170:
4169:
4165:
4164:
4161:
4160:
4148:
4142:
4140:
4134:
4132:
4129:
4128:
4125:
4124:
4120:
4119:
4116:
4115:
4107:Moselopteridae
4103:
4097:
4095:
4089:
4087:
4086:
4079:
4078:
4075:
4074:
4070:
4069:
4066:
4065:
4061:
4060:
4057:
4056:
4052:
4051:
4048:
4047:
4043:
4042:
4039:
4038:
4034:
4033:
4030:
4029:
4017:
4014:
4013:
4010:
4009:
3997:
3992:
3990:
3987:
3986:
3983:
3982:
3970:
3964:
3962:
3961:
3954:
3953:
3950:
3949:
3945:
3944:
3941:
3940:
3932:Kokomopteridae
3928:
3925:
3924:
3921:
3920:
3908:
3903:
3901:
3900:
3891:
3889:
3886:
3885:
3882:
3881:
3877:
3876:
3873:
3872:
3860:
3857:
3856:
3853:
3852:
3840:
3835:
3833:
3832:
3823:
3821:
3818:
3817:
3814:
3813:
3809:
3808:
3805:
3804:
3792:
3786:
3784:
3778:
3776:
3775:
3766:
3764:
3763:
3754:
3725:Parastylonurus
3707:
3706:
3697:
3696:
3695:
3686:
3685:
3684:
3683:
3682:
3680:
3677:
3673:
3672:
3667:
3666:
3662:
3661:
3658:
3657:
3653:
3652:
3649:
3648:
3644:
3643:
3640:
3639:
3635:
3634:
3631:
3630:
3626:
3625:
3622:
3621:
3617:
3616:
3613:
3612:
3608:
3607:
3604:
3603:
3599:
3598:
3595:
3594:
3590:
3589:
3586:
3585:
3581:
3580:
3577:
3576:
3572:
3571:
3568:
3567:
3563:
3562:
3559:
3558:
3550:
3547:
3546:
3543:
3542:
3535:
3530:
3528:
3527:
3520:
3519:
3516:
3515:
3507:
3502:
3500:
3499:
3492:
3491:
3488:
3487:
3477:
3474:
3473:
3470:
3469:
3465:
3464:
3461:
3460:
3450:
3447:
3446:
3443:
3442:
3432:
3427:
3425:
3422:
3421:
3418:
3417:
3407:
3404:
3403:
3400:
3399:
3395:
3394:
3391:
3390:
3380:
3377:
3376:
3373:
3372:
3362:
3357:
3355:
3349:
3347:
3346:
3339:
3338:
3335:
3334:
3330:
3329:
3326:
3325:
3321:
3320:
3317:
3316:
3312:
3311:
3308:
3307:
3303:
3302:
3299:
3298:
3291:
3288:
3287:
3284:
3283:
3275:
3270:
3268:
3265:
3264:
3261:
3260:
3250:
3244:
3242:
3241:
3234:
3233:
3230:
3229:
3219:
3214:
3212:
3209:
3208:
3205:
3204:
3194:
3188:
3186:
3185:
3176:
3174:
3171:
3170:
3167:
3166:
3156:
3150:
3148:
3145:
3144:
3141:
3140:
3136:
3135:
3132:
3131:
3121:
3118:
3117:
3114:
3113:
3103:
3098:
3096:
3090:
3088:
3085:
3084:
3081:
3080:
3070:
3064:
3062:
3061:
3054:
3053:
3050:
3049:
3039:
3034:
3032:
3031:
3024:
3023:
3020:
3019:
3015:
3014:
3011:
3010:
3006:
3005:
3002:
3001:
2991:
2988:
2987:
2984:
2983:
2974:
2969:
2967:
2964:
2963:
2960:
2959:
2949:
2943:
2941:
2940:
2931:
2929:
2928:
2921:
2920:
2917:
2916:
2912:
2911:
2908:
2907:
2903:
2902:
2899:
2898:
2888:
2885:
2884:
2881:
2880:
2870:
2865:
2863:
2860:
2859:
2856:
2855:
2845:
2839:
2837:
2836:
2832: â
2827:
2825:
2822:
2821:
2818:
2817:
2813:
2812:
2809:
2808:
2798:
2795:
2794:
2791:
2790:
2782:
2779:
2778:
2775:
2774:
2764:
2758:
2756:
2755:
2751: â
2746:
2744:
2741:
2740:
2737:
2736:
2726:
2720:
2718:
2717:
2708:
2664:Parastylonurus
2589:
2588:
2579:
2578:
2577:
2568:
2567:
2566:
2565:
2564:
2562:
2561:Classification
2559:
2502:Anthraconectes
2460:Malcolm Laurie
2446:Henry Woodward
2355:together with
2294:Hibbertopterus
2267:and the genus
2210:James E. DeKay
2155:
2152:
2117:Adelophthalmus
2111:respectively.
2090:Adelophthalmus
2085:Adelophthalmus
2075:Adelophthalmus
2067:Adelophthalmus
2063:Adelophthalmus
2058:Adelophthalmus
2052:Adelophthalmus
2047:Adelophthalmus
2041:Hibbertopterus
2037:Adelophthalmus
2031:, such as the
2007:hibbertopterid
1998:
1995:
1921:Adelophthalmus
1911:
1908:
1774:
1771:
1676:Fezouata Biota
1630:
1627:
1625:
1622:
1518:D. bembycoides
1496:
1489:
1488:
1487:
1477:
1470:
1469:
1468:
1467:
1466:
1464:
1461:
1457:apex predatory
1355:
1352:
1336:Strobilopterus
1263:(not to scale)
1260:Strobilopterus
1243:
1240:
1147:pseudotracheae
1138:
1135:
1119:P. kosinkiorum
1101:Hibbertopterus
1031:Parastylonurus
1025:Hibbertopterus
1021:Hibbertopterus
1016:Hibbertopterus
981:Hibbertopterus
977:
970:
969:
968:
949:
942:
941:
940:
939:
938:
936:
933:
929:Hibbertopterus
925:Hibbertopterus
858:The family of
751:
748:
746:
743:
634:Pterygotioidea
567:. As in other
530:
527:
512:Adelophthalmus
363:that form the
348:
347:
346:
345:
336:
329:Gigantostraca
324:
323:
317:
316:
314:
313:
305:
292:Incertae sedis
289:
288:
287:
286:
285:
273:
260:
259:
255:
254:
239:
235:
234:
229:
222:
221:
216:
212:
211:
206:
202:
201:
196:
192:
191:
186:
182:
181:
168:
167:
155:housed at the
146:
145:
137:
136:
121:
116:
111:
106:
101:
96:
91:
86:
81:
76:
71:
66:
48:
26:
9:
6:
4:
3:
2:
8952:
8941:
8938:
8936:
8933:
8931:
8928:
8926:
8923:
8921:
8918:
8916:
8913:
8912:
8910:
8893:
8888:
8884:
8880:
8875:
8871:
8867:
8862:
8858:
8854:
8849:
8845:
8841:
8836:
8832:
8828:
8823:
8819:
8814:
8808:
8804:
8799:
8793:
8789:
8788:
8786:
8784:
8780:
8776:
8771:
8761:
8751:
8745:
8742:
8740:
8737:
8736:
8734:
8730:
8724:
8721:
8719:
8716:
8714:
8711:
8710:
8708:
8704:
8698:
8697:
8693:
8691:
8690:
8686:
8684:
8683:
8679:
8678:
8676:
8674:
8670:
8664:
8663:
8659:
8657:
8656:
8652:
8650:
8649:
8645:
8643:
8642:
8641:Pentecopterus
8638:
8636:
8635:
8631:
8629:
8628:
8624:
8622:
8621:
8617:
8615:
8614:
8613:Megalograptus
8610:
8608:
8607:
8606:Jaekelopterus
8603:
8601:
8600:
8596:
8594:
8593:
8589:
8587:
8586:
8582:
8580:
8579:
8578:Drepanopterus
8575:
8573:
8572:
8568:
8566:
8565:
8561:
8559:
8558:
8557:Brachyopterus
8554:
8552:
8551:
8547:
8546:
8544:
8540:
8534:
8533:South America
8531:
8529:
8526:
8524:
8523:North America
8521:
8519:
8516:
8514:
8511:
8509:
8506:
8505:
8503:
8501:
8497:
8491:
8488:
8486:
8485:Carboniferous
8483:
8481:
8478:
8476:
8473:
8471:
8468:
8467:
8465:
8463:
8462:Geochronology
8459:
8454:
8432:
8429:
8427:
8424:
8422:
8419:
8418:
8416:
8414:
8410:
8404:
8401:
8400:
8398:
8396:
8392:
8386:
8383:
8382:
8380:
8378:
8374:
8368:
8365:
8363:
8360:
8358:
8355:
8354:
8352:
8350:
8346:
8343:
8341:
8335:
8329:
8326:
8324:
8321:
8319:
8316:
8315:
8313:
8311:
8310:Eurypteroidea
8307:
8301:
8298:
8297:
8295:
8293:
8289:
8283:
8280:
8279:
8277:
8275:
8271:
8268:
8266:
8260:
8250:
8249:Mycteroptidae
8247:
8245:
8242:
8240:
8237:
8236:
8234:
8232:
8231:Mycteropoidea
8228:
8222:
8219:
8217:
8214:
8213:
8211:
8209:
8205:
8199:
8196:
8194:
8191:
8190:
8188:
8186:
8185:Stylonuroidea
8182:
8176:
8175:Rhenopteridae
8173:
8172:
8170:
8168:
8164:
8161:
8159:
8153:
8147:
8144:
8143:
8141:
8137:
8134:
8130:
8124:
8123:Sclerophorata
8120:
8117:
8115:
8111:
8108:
8106:
8102:
8099:
8097:
8093:
8090:
8088:
8087:Euchelicerata
8084:
8081:
8079:
8075:
8073:
8069:
8067:
8063:
8062:
8059:
8055:
8048:
8043:
8041:
8036:
8034:
8029:
8028:
8025:
8019:
8016:
8013:
8010:
8009:
7999:
7995:
7991:
7986:
7985:
7974:
7970:
7966:
7962:
7958:
7954:
7949:
7945:
7941:
7937:
7933:
7928:
7923:
7919:
7915:
7912:(7068): 576.
7911:
7907:
7906:
7901:
7896:
7892:
7888:
7883:
7878:
7873:
7868:
7864:
7860:
7856:
7851:
7847:
7843:
7838:
7833:
7829:
7825:
7824:Palaeontology
7821:
7816:
7812:
7808:
7804:
7800:
7795:
7790:
7786:
7782:
7778:
7774:
7770:
7766:
7762:
7757:
7753:
7749:
7745:
7741:
7737:
7733:
7729:
7725:
7721:
7717:
7712:
7708:
7704:
7700:
7696:
7692:
7688:
7684:
7680:
7676:
7672:
7671:Palaeontology
7667:
7663:
7659:
7655:
7651:
7647:
7643:
7639:
7634:
7627:
7623:
7619:
7615:
7611:
7604:
7602:
7595:
7591:
7587:
7583:
7579:
7575:
7571:
7567:
7563:
7558:
7554:
7550:
7546:
7542:
7538:
7534:
7530:
7526:
7521:
7517:
7513:
7506:
7504:
7497:
7487:on 2020-06-06
7483:
7479:
7475:
7471:
7467:
7463:
7459:
7455:
7451:
7447:
7443:
7436:
7434:
7427:
7423:
7419:
7415:
7414:
7408:
7401:
7397:
7393:
7389:
7385:
7378:
7373:
7369:
7365:
7361:
7357:
7353:
7349:
7342:
7337:
7333:
7329:
7325:
7321:
7317:
7313:
7309:
7305:
7300:
7296:
7292:
7288:
7284:
7280:
7276:
7272:
7268:
7263:
7259:
7255:
7251:
7247:
7243:
7239:
7235:
7231:
7226:
7222:
7218:
7214:
7210:
7206:
7202:
7198:
7194:
7190:
7186:
7185:
7179:
7175:
7171:
7167:
7162:
7158:
7152:
7148:
7147:
7141:
7137:
7133:
7128:
7123:
7119:
7115:
7111:
7107:
7103:
7098:
7094:
7090:
7086:
7082:
7078:
7073:
7069:
7065:
7061:
7057:
7053:
7048:
7044:
7040:
7035:
7030:
7025:
7020:
7016:
7012:
7008:
7004:
7003:
6998:
6993:
6989:
6988:
6982:
6978:
6974:
6969:
6964:
6959:
6954:
6950:
6946:
6942:
6938:
6937:
6932:
6927:
6922:
6917:
6913:
6909:
6905:
6900:
6896:
6892:
6888:
6884:
6879:
6874:
6870:
6866:
6862:
6858:
6857:
6852:
6847:
6843:
6839:
6835:
6831:
6826:
6821:
6817:
6813:
6809:
6805:
6801:
6796:
6792:
6788:
6784:
6780:
6776:
6772:
6768:
6764:
6760:
6756:
6755:Palaeontology
6752:
6747:
6743:
6739:
6734:
6729:
6725:
6721:
6717:
6713:
6709:
6704:
6700:
6696:
6692:
6688:
6683:
6679:
6675:
6671:
6666:
6662:
6658:
6654:
6650:
6645:
6641:
6637:
6633:
6629:
6624:
6620:
6616:
6612:
6608:
6604:
6600:
6596:
6592:
6587:
6583:
6579:
6575:
6571:
6567:
6563:
6559:
6557:
6550:
6546:
6540:
6536:
6535:
6529:
6525:
6523:82-00-04963-9
6519:
6515:
6511:
6507:
6503:
6496:
6491:
6487:
6481:
6477:
6476:
6470:
6466:
6459:
6454:
6450:
6444:
6440:
6439:
6433:
6429:
6428:
6422:
6418:
6414:
6410:
6405:
6401:
6397:
6393:
6389:
6385:
6381:
6377:
6372:
6368:
6364:
6359:
6354:
6350:
6346:
6342:
6338:
6334:
6329:
6325:
6321:
6317:
6313:
6309:
6305:
6300:
6296:
6292:
6287:
6282:
6278:
6274:
6270:
6268:
6261:
6257:
6251:
6247:
6246:
6240:
6239:
6227:
6222:
6216:, p. 56.
6215:
6210:
6203:
6198:
6191:
6186:
6180:, p. 25.
6179:
6174:
6167:
6162:
6155:
6150:
6143:
6138:
6131:
6126:
6119:
6114:
6107:
6106:Lamsdell 2012
6102:
6096:, p. 19.
6095:
6094:Lamsdell 2012
6090:
6084:, p. 17.
6083:
6078:
6071:
6066:
6059:
6058:Kingsley 1894
6054:
6047:
6042:
6035:
6030:
6028:
6020:
6015:
6008:
6003:
5996:
5991:
5985:, p. 19.
5984:
5979:
5972:
5971:Woodward 1865
5967:
5961:, p. 14.
5960:
5955:
5949:, p. 18.
5948:
5943:
5936:
5931:
5924:
5919:
5913:, p. 27.
5912:
5907:
5900:
5895:
5888:
5883:
5877:, p. 13.
5876:
5871:
5864:
5859:
5852:
5847:
5845:
5837:
5832:
5826:, p. 19.
5825:
5820:
5813:
5808:
5801:
5796:
5790:, p. 24.
5789:
5784:
5778:, p. 79.
5777:
5772:
5765:
5760:
5752:
5748:
5744:
5740:
5736:
5732:
5728:
5724:
5720:
5716:
5712:
5705:
5697:
5693:
5689:
5685:
5681:
5677:
5673:
5669:
5665:
5661:
5657:
5650:
5643:
5638:
5631:
5626:
5619:
5614:
5612:
5604:
5599:
5593:, p. 70.
5592:
5587:
5585:
5583:
5575:
5570:
5563:
5558:
5551:
5546:
5539:
5534:
5528:, p. 44.
5527:
5522:
5515:
5510:
5503:
5498:
5496:
5494:
5492:
5490:
5482:
5477:
5470:
5465:
5459:, p. 15.
5458:
5453:
5447:, p. 13.
5446:
5441:
5435:, p. 12.
5434:
5429:
5423:, p. 29.
5422:
5417:
5415:
5413:
5405:
5400:
5393:
5388:
5386:
5379:, p. 11.
5378:
5373:
5371:
5363:
5358:
5351:
5346:
5339:
5338:Lamsdell 2014
5334:
5327:
5322:
5315:
5310:
5303:
5298:
5291:
5286:
5279:
5274:
5267:
5262:
5256:, p. 77.
5255:
5250:
5244:, p. 47.
5243:
5238:
5232:, p. 46.
5231:
5226:
5224:
5222:
5215:, p. 34.
5214:
5209:
5207:
5200:, p. 44.
5199:
5194:
5188:, p. 33.
5187:
5182:
5176:, p. 32.
5175:
5170:
5168:
5160:
5155:
5153:
5145:
5140:
5133:
5128:
5121:
5116:
5109:
5104:
5097:
5092:
5085:
5080:
5074:, p. 39.
5073:
5068:
5061:
5056:
5054:
5046:
5041:
5034:
5029:
5022:
5017:
5011:, p. 45.
5010:
5005:
4999:, p. 43.
4998:
4993:
4986:
4981:
4974:
4969:
4967:
4965:
4963:
4955:
4950:
4948:
4946:
4944:
4937:, p. 19.
4936:
4931:
4924:
4919:
4913:, p. 15.
4912:
4907:
4900:
4895:
4888:
4883:
4876:
4872:
4867:
4865:
4863:
4855:
4850:
4848:
4846:
4844:
4836:
4831:
4824:
4819:
4817:
4815:
4806:
4800:
4793:
4788:
4782:, p. 22.
4781:
4776:
4774:
4766:
4761:
4759:
4751:
4746:
4744:
4742:
4740:
4738:
4736:
4734:
4732:
4730:
4723:, p. 23.
4722:
4717:
4713:
4699:
4696:
4693:
4690:
4687:
4684:
4682:
4679:
4678:
4674:
4663:
4660:
4654:
4649:
4637:
4636:
4628:
4627:
4619:
4618:
4610:
4609:
4601:
4600:
4592:
4591:
4583:
4582:
4574:
4573:
4565:
4564:
4556:
4555:
4552:
4550:
4546:
4540:
4539:
4536:
4535:
4532:
4530:
4526:
4520:
4519:
4513:
4512:
4509:
4508:
4505:
4503:
4499:
4492:
4491:
4484:
4481:
4480:
4479:
4476:
4475:
4472:
4470:
4466:
4460:
4459:
4453:
4452:
4449:
4448:
4440:
4439:
4436:
4434:
4430:
4423:
4422:
4415:
4414:
4408:
4407:
4404:
4403:
4395:
4394:
4386:
4385:
4382:
4380:
4376:
4370:
4369:
4366:
4365:
4362:
4360:
4356:
4350:
4349:
4343:
4342:
4339:
4338:
4330:
4329:
4326:
4324:
4320:
4313:
4312:
4305:
4304:
4297:
4294:
4293:
4292:
4285:
4282:
4281:
4280:
4277:
4276:
4268:
4267:
4259:
4258:
4255:
4253:
4249:
4243:
4242:
4239:
4238:
4235:
4233:
4229:
4223:
4222:
4216:
4215:
4212:
4211:
4208:
4206:
4202:
4195:
4194:
4187:
4186:Eurypteroidea
4184:
4183:
4182:
4176:
4175:
4172:
4171:
4163:
4162:
4159:
4157:
4153:
4146:
4145:
4138:
4137:
4131:
4130:
4127:
4126:
4118:
4117:
4114:
4112:
4108:
4101:
4100:
4093:
4092:
4085:
4082:
4081:
4080:
4077:
4076:
4068:
4067:
4059:
4058:
4050:
4049:
4041:
4040:
4032:
4031:
4028:
4026:
4022:
4021:Mycteroptidae
4016:
4015:
4012:
4011:
4008:
4006:
4002:
3996:
3995:
3989:
3988:
3985:
3984:
3981:
3979:
3975:
3968:
3967:
3960:
3959:Mycteropoidea
3957:
3956:
3955:
3952:
3951:
3943:
3942:
3939:
3937:
3933:
3927:
3926:
3923:
3922:
3919:
3917:
3913:
3907:
3906:
3899:
3896:
3895:
3894:
3888:
3887:
3884:
3883:
3875:
3874:
3871:
3869:
3865:
3859:
3858:
3855:
3854:
3851:
3849:
3845:
3839:
3838:
3831:
3830:Stylonuroidea
3828:
3827:
3826:
3820:
3819:
3816:
3815:
3807:
3806:
3803:
3801:
3797:
3796:Rhenopteridae
3790:
3789:
3782:
3781:
3774:
3771:
3770:
3769:
3762:
3759:
3758:
3757:
3753:
3750:
3744:
3740:
3736:
3727:
3726:
3721:
3717:
3716:
3711:
3701:
3690:
3669:
3668:
3660:
3659:
3651:
3650:
3642:
3641:
3633:
3632:
3624:
3623:
3615:
3614:
3606:
3605:
3597:
3596:
3588:
3587:
3579:
3578:
3570:
3569:
3561:
3560:
3557:
3556:
3549:
3548:
3545:
3544:
3541:
3540:
3534:
3533:
3526:
3525:Sclerophorata
3523:
3522:
3521:
3518:
3517:
3514:
3513:
3506:
3505:
3498:
3495:
3494:
3493:
3490:
3489:
3486:
3485:
3484:
3476:
3475:
3472:
3471:
3463:
3462:
3459:
3458:
3457:
3449:
3448:
3445:
3444:
3441:
3440:
3439:
3431:
3430:
3424:
3423:
3420:
3419:
3416:
3415:
3414:
3413:Pasternakevia
3406:
3405:
3402:
3401:
3393:
3392:
3389:
3388:
3387:
3386:Cyamocephalus
3379:
3378:
3375:
3374:
3371:
3370:
3369:
3361:
3360:
3353:
3352:
3345:
3342:
3341:
3340:
3337:
3336:
3328:
3327:
3319:
3318:
3310:
3309:
3301:
3300:
3297:
3296:
3290:
3289:
3286:
3285:
3282:
3281:
3274:
3273:
3267:
3266:
3263:
3262:
3259:
3258:
3257:
3248:
3247:
3240:
3237:
3236:
3235:
3232:
3231:
3228:
3227:
3226:
3218:
3217:
3211:
3210:
3207:
3206:
3203:
3202:
3201:
3200:Kasibelinurus
3192:
3191:
3184:
3181:
3180:
3179:
3173:
3172:
3169:
3168:
3165:
3164:
3163:
3154:
3153:
3147:
3146:
3143:
3142:
3134:
3133:
3130:
3129:
3128:
3120:
3119:
3116:
3115:
3112:
3111:
3110:
3102:
3101:
3094:
3093:
3087:
3086:
3083:
3082:
3079:
3078:
3077:
3068:
3067:
3060:
3057:
3056:
3055:
3052:
3051:
3048:
3047:
3046:
3038:
3037:
3030:
3029:Euchelicerata
3027:
3026:
3025:
3022:
3021:
3013:
3012:
3004:
3003:
3000:
2999:
2998:
2990:
2989:
2986:
2985:
2982:
2981:
2980:
2973:
2972:
2966:
2965:
2962:
2961:
2958:
2957:
2956:
2947:
2946:
2939:
2936:
2935:
2934:
2927:
2924:
2923:
2922:
2919:
2918:
2910:
2909:
2901:
2900:
2897:
2896:
2895:
2887:
2886:
2883:
2882:
2879:
2878:
2877:
2876:Alalcomenaeus
2869:
2868:
2862:
2861:
2858:
2857:
2854:
2853:
2852:
2843:
2842:
2835:
2831:
2830:
2824:
2823:
2820:
2819:
2811:
2810:
2807:
2806:
2805:
2797:
2796:
2793:
2792:
2789:
2788:
2781:
2780:
2777:
2776:
2773:
2772:
2771:
2762:
2761:
2754:
2750:
2749:
2743:
2742:
2739:
2738:
2735:
2734:
2733:
2724:
2723:
2716:
2713:
2712:
2711:
2707:
2705:
2701:
2697:
2693:
2689:
2688:Sclerophorata
2685:
2681:
2677:
2676:
2671:
2670:
2665:
2661:
2656:
2655:Ernst Haeckel
2651:
2649:
2648:Ray Lankester
2645:
2640:
2635:
2633:
2628:
2611:
2610:Sclerophorata
2607:
2606:
2605:Heterophrynus
2601:
2597:
2593:
2583:
2572:
2558:
2556:
2552:
2548:
2547:
2542:
2538:
2534:
2530:
2529:Drepanopterus
2526:
2525:
2520:
2519:
2518:Dolichopterus
2514:
2513:
2508:
2504:
2503:
2498:
2493:
2489:
2485:
2480:
2478:
2474:
2470:
2466:
2461:
2457:
2453:
2452:
2447:
2442:
2440:
2436:
2432:
2428:
2424:
2420:
2416:
2412:
2408:
2400:
2396:
2392:
2388:
2384:
2382:
2378:
2374:
2373:
2368:
2364:
2363:
2358:
2354:
2350:
2345:
2343:
2339:
2335:
2331:
2327:
2326:Louis Agassiz
2307:
2303:
2299:
2295:
2290:
2288:
2285:(composed of
2284:
2280:
2276:
2272:
2271:
2266:
2262:
2260:
2254:
2249:
2246:
2240:
2234:
2228:
2223:
2219:
2215:
2211:
2207:
2203:
2202:
2197:
2196:Oneida county
2193:
2189:
2185:
2177:
2173:
2172:
2166:
2161:
2151:
2149:
2145:
2140:
2139:
2134:
2130:
2126:
2122:
2118:
2114:
2110:
2109:
2104:
2103:
2097:
2095:
2091:
2086:
2083:
2082:Early Permian
2078:
2076:
2072:
2068:
2064:
2059:
2055:
2053:
2049:
2048:
2042:
2038:
2034:
2030:
2021:
2017:
2013:
2012:
2008:
2003:
1994:
1990:
1988:
1982:
1979:
1975:
1970:
1966:
1961:
1959:
1955:
1951:
1947:
1946:Jaekelopterus
1939:
1935:
1934:Carboniferous
1931:
1927:
1923:
1922:
1916:
1907:
1905:
1901:
1897:
1893:
1887:
1885:
1880:
1876:
1872:
1868:
1864:
1860:
1856:
1852:
1848:
1844:
1840:
1836:
1831:
1829:
1825:
1824:Mycteropoidea
1821:
1817:
1816:Stylonuroidea
1813:
1807:
1805:
1804:Pridoli epoch
1798:
1794:
1790:
1786:
1785:
1779:
1770:
1768:
1767:Pentecopterus
1764:
1763:Megalograptus
1760:
1759:
1758:Echinognathus
1754:
1750:
1746:
1742:
1738:
1737:pseudofossils
1733:
1731:
1727:
1723:
1719:
1718:
1713:
1708:
1706:
1702:
1701:
1700:Brachyopterus
1695:
1694:Pentecopterus
1691:
1689:
1685:
1681:
1677:
1673:
1669:
1665:
1664:Pentecopterus
1662:
1661:megalograptid
1658:
1650:
1647:belongs, the
1646:
1645:Pentecopterus
1642:
1641:
1640:Pentecopterus
1635:
1621:
1611:
1601:
1593:
1588:
1584:
1582:
1577:
1576:spermatophore
1573:
1569:
1559:
1548:
1542:
1537:
1534:
1530:
1525:
1523:
1519:
1515:
1514:
1513:Drepanopterus
1502:
1501:
1493:
1483:
1482:
1474:
1460:
1458:
1453:
1451:
1450:Lanarkopterus
1447:
1443:
1442:
1437:
1433:
1428:
1426:
1422:
1421:
1420:Buffalopterus
1416:
1415:
1410:
1406:
1402:
1397:
1394:
1390:
1389:
1388:Megalograptus
1384:
1380:
1372:
1371:
1366:
1365:
1360:
1351:
1349:
1345:
1340:
1338:
1337:
1332:
1331:Jaekelopterus
1328:
1324:
1320:
1316:
1311:
1309:
1305:
1301:
1297:
1293:
1288:
1286:
1282:
1278:
1274:
1270:
1262:
1261:
1256:
1252:
1248:
1239:
1231:
1225:
1223:
1219:
1215:
1205:
1204:pseudotrachea
1201:
1197:
1192:
1188:
1182:
1180:
1179:invaginations
1176:
1162:
1161:
1156:
1152:
1148:
1143:
1134:
1132:
1131:
1126:
1125:
1120:
1116:
1115:
1110:
1106:
1102:
1097:
1095:
1091:
1090:
1081:
1080:
1075:
1070:
1066:
1064:
1060:
1056:
1052:
1048:
1044:
1043:water beetles
1040:
1039:Eurypteroidea
1035:
1033:
1032:
1026:
1022:
1017:
1013:
1008:
1006:
1002:
998:
987:
983:
982:
974:
963:
959:
955:
954:
946:
932:
930:
926:
922:
917:
915:
914:
909:
905:
900:
898:
897:
892:
891:
885:
882:
878:
876:
871:
867:
866:
861:
860:Jaekelopterus
856:
854:
850:
846:
845:
840:
839:Jaekelopterus
836:
835:
829:
828:Jaekelopterus
824:
820:
817:
813:
812:
806:
804:
803:
798:
791:
790:
785:
784:
779:
775:
774:
769:
768:
763:
762:
756:
742:
738:
733:
727:
725:
721:
710:
709:Kiemenplatten
704:
677:
671:
669:
665:
661:
657:
653:
652:
647:
643:
642:Mycteroptidae
639:
635:
631:
626:
624:
621:
617:
613:
609:
605:
601:
596:
594:
590:
589:compound eyes
586:
582:
578:
574:
570:
566:
562:
558:
554:
550:
542:
541:
535:
526:
524:
521:, achieved a
520:
519:
514:
513:
508:
504:
500:
499:North America
496:
495:
490:
489:
488:Jaekelopterus
483:
481:
479:
472:
467:
465:
458:
453:
452:Ancient Greek
449:
445:
441:
437:
433:
429:
424:
418:
414:
410:
406:
405:
400:
396:
393:
389:
388:Late Cambrian
385:
381:
378:period 467.3
377:
374:stage of the
373:
369:
366:
362:
358:
357:sea scorpions
354:
342:
338:Cyrtoctenida
337:
333:
328:
327:
325:
322:
318:
312:
311:
310:Marsupipterus
306:
304:
303:
298:
297:
293:
282:
278:
274:
268:
264:
263:
261:
256:
251:
246:
240:
237:
236:
233:
232:Sclerophorata
230:
227:
224:
223:
220:
217:
214:
213:
210:
207:
204:
203:
200:
197:
194:
193:
190:
187:
184:
183:
178:
173:
169:
166:
162:
158:
154:
153:
147:
143:
138:
132:
124:
119:
114:
109:
104:
99:
94:
89:
84:
79:
74:
69:
63:
56:
52:
44:
41:
37:
33:
19:
8782:
8694:
8687:
8680:
8660:
8653:
8646:
8639:
8632:
8625:
8618:
8611:
8604:
8599:Hughmilleria
8597:
8590:
8583:
8576:
8569:
8562:
8555:
8548:
8431:Pterygotidae
8367:Mixopteridae
8323:Eurypteridae
8198:Stylonuridae
8118:
8109:
8100:
8091:
8082:
8053:
7998:the original
7993:
7956:
7952:
7909:
7903:
7862:
7858:
7827:
7823:
7768:
7764:
7719:
7715:
7674:
7670:
7648:(1): 19â30.
7645:
7641:
7637:
7626:the original
7613:
7609:
7600:
7565:
7561:
7528:
7524:
7515:
7511:
7502:
7489:. Retrieved
7482:the original
7445:
7441:
7432:
7412:
7400:the original
7387:
7383:
7351:
7347:
7315:
7311:
7308:Vinetopterus
7307:
7304:Alkenopterus
7303:
7270:
7266:
7236:(1): 13â27.
7233:
7229:
7191:(1): 15â45.
7188:
7182:
7173:
7169:
7145:
7109:
7105:
7084:
7080:
7059:
7055:
7009:(169): 169.
7006:
7000:
6986:
6940:
6934:
6911:
6907:
6860:
6854:
6810:(1): 49â61.
6807:
6803:
6758:
6754:
6750:
6715:
6711:
6690:
6686:
6677:
6673:
6652:
6648:
6631:
6627:
6594:
6590:
6568:(1): 15â18.
6565:
6561:
6555:
6533:
6505:
6501:
6474:
6464:
6437:
6426:
6416:
6412:
6386:(1): 39â45.
6383:
6379:
6375:
6340:
6336:
6307:
6303:
6276:
6272:
6266:
6244:
6235:Bibliography
6228:, p. 3.
6221:
6209:
6197:
6185:
6173:
6161:
6149:
6137:
6125:
6113:
6101:
6089:
6077:
6065:
6053:
6041:
6014:
6002:
5990:
5978:
5966:
5954:
5942:
5930:
5918:
5906:
5894:
5882:
5870:
5858:
5831:
5819:
5807:
5802:, p. 1.
5795:
5783:
5771:
5759:
5718:
5714:
5704:
5663:
5659:
5649:
5637:
5625:
5598:
5569:
5564:, p. 3.
5557:
5545:
5533:
5521:
5509:
5476:
5464:
5452:
5440:
5428:
5406:, p. 6.
5399:
5394:, p. 1.
5357:
5345:
5333:
5321:
5309:
5297:
5285:
5273:
5261:
5249:
5237:
5193:
5181:
5139:
5127:
5115:
5103:
5091:
5079:
5067:
5040:
5028:
5016:
5004:
4992:
4980:
4930:
4918:
4906:
4894:
4882:
4830:
4799:
4787:
4721:Størmer 1955
4716:
4543:
4525:Pterygotidae
4523:
4496:
4463:
4427:
4375:Mixopteridae
4373:
4353:
4317:
4246:
4228:Eurypteridae
4226:
4199:
4150:
4105:
4019:
3999:
3972:
3930:
3910:
3864:Stylonuridae
3862:
3842:
3794:
3760:
3747:between the
3745:
3741:
3737:
3733:
3723:
3713:
3554:
3552:
3537:
3509:
3481:
3479:
3454:
3452:
3436:
3434:
3411:
3409:
3384:
3382:
3368:Pseudoniscus
3366:
3364:
3293:
3277:
3254:
3252:
3225:Willwerathia
3223:
3221:
3198:
3196:
3160:
3158:
3125:
3123:
3107:
3105:
3074:
3072:
3043:
3041:
2995:
2993:
2977:
2976:
2955:Palaeoisopus
2953:
2951:
2892:
2890:
2874:
2872:
2849:
2847:
2802:
2800:
2784:
2768:
2766:
2730:
2728:
2680:paraphyletic
2673:
2667:
2663:
2659:
2652:
2636:
2626:
2617:
2603:
2594:such as the
2550:
2546:Hughmilleria
2544:
2540:
2537:Erettopterus
2536:
2532:
2528:
2524:Onychopterus
2522:
2516:
2510:
2506:
2500:
2496:
2491:
2481:
2468:
2464:
2455:
2449:
2443:
2434:
2422:
2418:
2414:
2410:
2406:
2404:
2376:
2370:
2367:Entomostraca
2360:
2356:
2352:
2346:
2333:
2329:
2305:
2297:
2296:in 1836 and
2293:
2291:
2286:
2283:Eurypteridae
2274:
2268:
2264:
2256:
2250:
2221:
2218:Branchiopoda
2213:
2199:
2192:Westmoreland
2181:
2169:
2136:
2125:C. permianus
2124:
2120:
2116:
2112:
2106:
2100:
2098:
2089:
2084:
2079:
2074:
2066:
2062:
2057:
2056:
2051:
2045:
2040:
2036:
2025:
2015:
2009:
1991:
1983:
1962:
1945:
1943:
1919:
1899:
1891:
1888:
1883:
1866:
1858:
1854:
1834:
1832:
1808:
1801:
1797:Pterygotidae
1784:Erettopterus
1782:
1766:
1762:
1756:
1734:
1715:
1709:
1698:
1693:
1692:
1690:appendages.
1663:
1654:
1644:
1638:
1612:
1597:
1591:
1568:spermathecae
1540:
1538:
1532:
1528:
1526:
1521:
1517:
1511:
1508:
1498:
1479:
1454:
1449:
1446:jawless fish
1439:
1431:
1429:
1418:
1412:
1408:
1404:
1398:
1386:
1382:
1375:
1368:
1362:
1343:
1341:
1334:
1330:
1312:
1296:megacheirans
1289:
1266:
1258:
1230:asphyxiation
1226:
1193:
1187:Kiemenplatte
1183:
1166:
1158:
1128:
1122:
1118:
1112:
1105:South Africa
1100:
1098:
1087:
1085:
1077:
1036:
1029:
1024:
1020:
1015:
1009:
994:
979:
951:
928:
924:
920:
918:
913:Arthropleura
911:
901:
894:
888:
886:
880:
873:
870:A. bohemicus
869:
863:
859:
857:
852:
848:
842:
838:
832:
827:
809:
807:
800:
794:
787:
781:
778:A. bohemicus
777:
771:
765:
759:
728:
672:
649:
627:
608:Pterygotidae
597:
569:chelicerates
559:composed of
546:
538:
516:
510:
494:Alkenopterus
492:
486:
484:
477:
475:
463:
461:
447:
425:
402:
367:
356:
352:
351:
308:
300:
291:
244:
225:
150:
131:Jiangshanian
55:Late Permian
40:
36:Eurypteridae
34:itself, see
8920:Eurypterida
8848:iNaturalist
8813:Eurypterida
8807:Wikispecies
8783:Eurypterida
8696:Palmichnium
8673:Ichnogenera
8571:Carcinosoma
8265:Eurypterina
8158:Stylonurina
8096:Prosomapoda
8078:Chelicerata
8076:Subphylum:
8054:Eurypterids
7176:(3): 1â278.
6508:: 255â270.
6202:Tetlie 2007
5642:Tetlie 2007
5574:Tetlie 2007
5538:Tetlie 2006
5526:Selden 1999
5502:Tetlie 2007
5481:Tetlie 2007
5469:Tetlie 2007
5242:Selden 1999
5230:Selden 1999
5159:Selden 1985
5144:Selden 1985
5132:Selden 1985
5120:Selden 1985
5108:Selden 1985
5021:Selden 1999
5009:Selden 1999
4997:Selden 1999
4954:Tetlie 2007
4935:Tetlie 2008
4887:Briggs 1985
4823:Tetlie 2007
4084:Eurypterina
3773:Stylonurina
3761:Eurypterida
3720:Stylonurina
3715:Bassipterus
3710:Eurypterina
3555:Eurypterida
3483:Bembicosoma
3239:Xiphosurida
3162:Legrandella
3076:Weinbergina
3059:Prosomapoda
2938:Pycnogonida
2926:Chelicerata
2894:Leanchoilia
2770:Emeraldella
2753:Antennulata
2704:Xiphosurida
2700:Prosomapoda
2669:Rhenopterus
2622:BlattfĂźssen
2592:xiphosurans
2469:E. fischeri
2351:classified
2119:had, while
1954:brachiopods
1926:eurypterine
1924:, the only
1726:aglaspidids
1712:Precambrian
1680:Tremadocian
1668:Darriwilian
1436:cannibalism
1405:Carcinosoma
1300:crustaceans
1285:sea spiders
1281:xiphosurans
1235:BlattfĂźssen
1209:BlattfĂźssen
1184:Though the
1163:(pictured).
1137:Respiration
1114:Palmichnium
1055:sea turtles
997:Stylonurina
986:stylonurine
962:eurypterine
908:exoskeleton
715:BlattfĂźssen
620:Eurypterina
581:opisthosoma
529:Description
448:Eurypterida
436:fresh water
395:chelicerate
372:Darriwilian
368:Eurypterida
353:Eurypterids
302:Dorfopterus
277:Stylonurina
267:Eurypterina
245:Eurypterida
219:Chelicerata
215:Subphylum:
51:Darriwilian
18:Eurypterida
8909:Categories
8662:Stylonurus
8648:Pterygotus
8627:Mixopterus
8620:Megarachne
8585:Eurypterus
8470:Ordovician
8426:Slimonidae
8337:Infraorder
8114:Dekatriata
8105:Planaterga
8072:Arthropoda
7601:Eurypterus
7491:2018-10-09
7433:Eurypterus
7422:B0043KRIVC
7184:Cladistics
6943:(98): 98.
6419:: 157â158.
6248:. Norton.
5935:M'Coy 1849
5836:White 1927
4985:Whyte 2005
4704:References
4545:Slimonidae
3722:, such as
3712:, such as
3497:Dekatriata
3456:Limuloides
3344:Planaterga
3280:Belinurina
3256:Lunataspis
3109:Venustulus
2979:Pycnogonum
2834:Megacheira
2732:Fuxianhuia
2715:Arthropoda
2692:Dekatriata
2660:Eurypterus
2627:Pterygotus
2533:Pterygotus
2507:Stylonurus
2497:Eurypterus
2490:published
2477:homologies
2451:Stylonurus
2439:James Hall
2437:(1859) by
2415:Eurypterus
2381:David Page
2377:Pterygotus
2357:Eurypterus
2353:Pterygotus
2334:Eurypterus
2330:Pterygotus
2319:winged one
2306:Pterygotus
2287:Eurypterus
2281:) and the
2265:Eurypterus
2259:Trilobiten
2222:Eurypterus
2168:Figure of
2158:See also:
2094:Leonardian
2033:rhizodonts
1987:placoderms
1900:Pterygotus
1892:Acutiramus
1884:Eurypterus
1875:equatorial
1871:Euramerica
1867:Eurypterus
1859:Eurypterus
1855:E. remipes
1847:scavenging
1839:generalist
1835:Eurypterus
1730:strabopids
1657:Ordovician
1610:material.
1592:Eurypterus
1581:ovipositor
1563:BlattfĂźsse
1541:Eurypterus
1522:D. lobatus
1414:Eurypterus
1409:Acutiramus
1383:Mixopterus
1364:Pterygotus
1344:Eurypterus
1302:and basal
1292:trilobites
1170:BlattfĂźsse
1157:, such as
1089:Mixopterus
984:, a large
953:Eurypterus
935:Locomotion
875:Pterygotus
865:Acutiramus
849:E. grandis
698:BlattfĂźsse
676:BlattfĂźsse
648:, notably
604:homologous
600:chelicerae
549:arthropods
540:Eurypterus
518:Pterygotus
507:Euramerica
404:Eurypterus
376:Ordovician
361:arthropods
258:Suborders
250:Burmeister
209:Arthropoda
46:Eurypterid
32:Eurypterus
8739:Metastoma
8723:Xiphosura
8713:Arachnida
8500:Geography
8064:Kingdom:
7973:129427356
7846:133765946
7811:129319753
7803:0016-7649
7699:0024-4082
7622:0366-0176
7553:0031-0182
7470:1478-0941
7396:0038-6804
7390:: 39â54.
7332:127313845
7295:252467508
7287:0891-2963
7258:1502-3931
7205:1502-3931
6895:221590821
6834:1478-0941
6791:129243775
6783:1475-4983
6582:131539776
6295:0024-4082
6245:Evolution
5751:246819305
5743:0891-2963
5688:1477-2019
4709:Citations
3539:Arachnida
3183:Xiphosura
3127:Camanchia
3045:Offacolus
2997:Haliestes
2653:In 1866,
2600:arachnids
2482:In 1912,
2444:In 1865,
2362:Belinurus
2347:In 1849,
2342:Seraphims
2328:in 1839.
2275:Cytherina
2270:Cytherina
2251:In 1843,
2248:'wing').
2129:Kungurian
2113:Hastimima
2102:Hastimima
1969:Famennian
1843:predation
1741:Laurentia
1533:operculum
1529:metastoma
1308:arachnids
1304:myriapods
1151:posterior
1059:sea lions
1001:sea floor
690:leaf-feet
656:scorpions
651:Eusarcana
553:segmented
440:scorpions
392:Paleozoic
195:Kingdom:
189:Eukaryota
161:Karlsruhe
129:Possible
8822:BugGuide
8792:Wikidata
8760:Category
8682:Arcuites
8655:Slimonia
8480:Devonian
8475:Silurian
8262:Suborder
8155:Suborder
8132:Taxonomy
8070:Phylum:
8066:Animalia
7982:Websites
7936:16319874
7891:16587225
7752:46953627
7707:84268058
7662:85862868
7590:88413267
7518:: 79â90.
7478:83519549
7221:85744103
7213:34814371
7136:26289442
7043:26324341
6977:23663507
6914:: 1â27.
6887:32916114
6842:85398946
6742:19828493
6619:83826460
6400:90662577
6367:18029297
5696:59488956
4692:Cottidae
4645:See also
4493:
4416:
4306:
4196:
4139:
4094:
3969:
3783:
3354:
3295:Limulina
3249:
3193:
3155:
3095:
3069:
2948:
2844:
2804:Sidneyia
2763:
2725:
2639:subclass
2602:such as
2551:Strabops
2541:Slimonia
2512:Eusarcus
2458:(1893),
2401:in 1912.
2372:Slimonia
2338:Scotland
2279:ostracod
2184:New York
1965:Frasnian
1940:periods.
1910:Devonian
1793:Devonian
1789:Silurian
1773:Silurian
1753:Avalonia
1749:seafloor
1745:demersal
1722:Missouri
1717:Strabops
1707:period.
1705:Cambrian
1688:peytoiid
1678:of Late
1452:itself.
1370:Birkenia
1298:, basal
1277:ontogeny
1242:Ontogeny
1222:pleopods
1218:tracheae
1200:plastron
1175:spinules
1130:Arcuites
1109:Gondwana
1074:holotype
732:anterior
701:and the
668:metasoma
664:mesosoma
640:and the
623:suborder
591:and the
585:carapace
561:proteins
432:brackish
409:Devonian
399:Silurian
321:Synonyms
205:Phylum:
199:Animalia
185:Domain:
8879:5308281
8840:4305531
8528:Oceania
8490:Permian
7944:4422644
7914:Bibcode
7882:1085121
7773:Bibcode
7744:1305624
7724:Bibcode
7679:Bibcode
7570:Bibcode
7533:Bibcode
7450:Bibcode
7356:Bibcode
7238:Bibcode
7230:Lethaia
7127:4571687
7034:4556007
7011:Bibcode
6968:3679797
6945:Bibcode
6865:Bibcode
6812:Bibcode
6763:Bibcode
6733:2865068
6699:1304420
6680:: 5â50.
6661:1301554
6640:1301214
6611:2452113
6358:2412931
6312:Bibcode
5723:Bibcode
5668:Bibcode
4973:Palaeos
4686:Nepidae
4314:
3671:
3438:Bunodes
2313:
2206:catfish
2201:Silurus
2178:(1825).
2133:Roadian
2071:Pangaea
1938:Permian
1879:Baltica
1684:Morocco
1629:Origins
1455:Though
1393:derived
1354:Feeding
1327:Wyoming
1315:Pragian
1273:ecdysis
1269:instars
1255:instars
1214:isopods
1160:Oniscus
1155:isopods
904:molting
823:Germany
745:Biology
703:ventral
684:
577:prosoma
573:tagmata
557:cuticle
341:Størmer
332:Haeckel
238:Order:
165:Germany
8853:317819
8827:505839
8798:Q19436
8518:Europe
8508:Africa
8139:Genera
7971:
7942:
7934:
7905:Nature
7889:
7879:
7844:
7809:
7801:
7750:
7742:
7705:
7697:
7660:
7620:
7588:
7551:
7476:
7468:
7420:
7394:
7330:
7293:
7285:
7256:
7219:
7211:
7203:
7153:
7134:
7124:
7041:
7031:
6975:
6965:
6893:
6885:
6840:
6832:
6789:
6781:
6740:
6730:
6697:
6659:
6638:
6617:
6609:
6580:
6541:
6520:
6482:
6445:
6398:
6365:
6355:
6293:
6252:
5749:
5741:
5694:
5686:
2851:Yohoia
2245:ĎĎÎľĎĎν
2239:pteron
2027:gone,
1812:strata
1743:, and
1558:dorsal
1391:). In
1317:-aged
1251:Larval
847:, but
841:) and
816:Emsian
786:, and
737:Zipfel
636:, the
630:telson
612:chelae
593:ocelli
565:chitin
503:Europe
478:pterĂłn
471:ĎĎÎľĎĎν
454:words
428:marine
421:
334:, 1866
283:, 1924
281:Diener
252:, 1843
133:record
8892:18947
8866:11995
8861:IRMNG
8119:Clade
8110:Clade
8101:Clade
8092:Clade
8083:Clade
7969:S2CID
7940:S2CID
7842:S2CID
7807:S2CID
7748:S2CID
7740:JSTOR
7703:S2CID
7658:S2CID
7629:(PDF)
7606:(PDF)
7586:S2CID
7508:(PDF)
7485:(PDF)
7474:S2CID
7438:(PDF)
7403:(PDF)
7380:(PDF)
7344:(PDF)
7328:S2CID
7291:S2CID
7217:S2CID
6891:S2CID
6838:S2CID
6787:S2CID
6695:JSTOR
6657:JSTOR
6636:JSTOR
6615:S2CID
6607:JSTOR
6578:S2CID
6498:(PDF)
6461:(PDF)
6396:S2CID
5747:S2CID
5692:S2CID
2555:class
2233:Îľá˝ĎĎĎ
2227:eurĂşs
1851:worms
1617:furca
1606:furca
1572:sperm
1553:furca
1547:furca
1202:or a
1005:crabs
660:venom
616:coxae
464:eurĂşs
457:Îľá˝ĎĎĎ
365:order
226:Clade
8513:Asia
7932:PMID
7887:PMID
7799:ISSN
7695:ISSN
7618:ISSN
7549:ISSN
7466:ISSN
7418:ASIN
7392:ISSN
7306:and
7283:ISSN
7254:ISSN
7209:PMID
7201:ISSN
7151:ISBN
7132:PMID
7039:PMID
6973:PMID
6883:PMID
6830:ISSN
6779:ISSN
6738:PMID
6539:ISBN
6518:ISBN
6480:ISBN
6443:ISBN
6363:PMID
6291:ISSN
6250:ISBN
5739:ISSN
5684:ISSN
2672:and
2543:and
2527:and
2486:and
2429:and
2397:and
2359:and
2310:lit.
2105:and
1958:reef
1956:and
1936:and
1837:, a
1822:and
1791:and
1765:and
1520:and
1425:anus
1411:and
1385:and
1333:and
1283:and
1196:lung
1072:The
1057:and
1051:lift
958:lift
750:Size
681:lit.
563:and
515:and
501:and
68:Preę
8835:EoL
7961:doi
7922:doi
7910:438
7877:PMC
7867:doi
7832:doi
7789:hdl
7781:doi
7769:172
7732:doi
7687:doi
7650:doi
7614:118
7578:doi
7541:doi
7529:252
7458:doi
7364:doi
7352:309
7320:doi
7275:doi
7246:doi
7193:doi
7122:PMC
7114:doi
7089:doi
7064:doi
7029:PMC
7019:doi
6963:PMC
6953:doi
6916:doi
6912:167
6873:doi
6820:doi
6771:doi
6728:PMC
6720:doi
6599:doi
6570:doi
6510:doi
6388:doi
6353:PMC
6345:doi
6320:doi
6281:doi
6277:120
5731:doi
5676:doi
2425:by
2194:in
2174:by
1845:or
1728:or
1594:sp.
1524:).
1321:in
1257:of
1076:of
1007:).
434:or
386:or
159:in
8911::
8889::
8876::
8863::
8850::
8837::
8824::
8809::
8794::
8121::
8112::
8103::
8094::
8085::
7992:.
7967:.
7957:21
7955:.
7938:.
7930:.
7920:.
7908:.
7902:.
7885:.
7875:.
7863:13
7861:.
7857:.
7840:.
7828:61
7826:.
7822:.
7805:.
7797:.
7787:.
7779:.
7767:.
7763:.
7746:.
7738:.
7730:.
7720:63
7718:.
7701:.
7693:.
7685:.
7675:52
7673:.
7656:.
7646:49
7644:.
7612:.
7608:.
7584:.
7576:.
7564:.
7547:.
7539:.
7527:.
7516:76
7514:.
7510:.
7472:.
7464:.
7456:.
7444:.
7440:.
7388:32
7386:.
7382:.
7362:.
7350:.
7346:.
7326:.
7316:84
7314:.
7289:.
7281:.
7271:34
7269:.
7252:.
7244:.
7234:21
7232:.
7215:.
7207:.
7199:.
7189:29
7187:.
7174:11
7172:.
7168:.
7130:.
7120:.
7110:11
7108:.
7104:.
7083:.
7079:.
7060:17
7058:.
7054:.
7037:.
7027:.
7017:.
7007:15
7005:.
6999:.
6971:.
6961:.
6951:.
6941:13
6939:.
6933:.
6910:.
6906:.
6889:.
6881:.
6871:.
6861:30
6859:.
6853:.
6836:.
6828:.
6818:.
6806:.
6802:.
6785:.
6777:.
6769:.
6759:52
6757:.
6736:.
6726:.
6714:.
6710:.
6691:55
6689:.
6678:40
6676:.
6672:.
6653:38
6651:.
6632:35
6630:.
6613:.
6605:.
6595:28
6593:.
6576:.
6564:.
6560:.
6516:.
6504:.
6500:.
6463:.
6417:33
6415:.
6411:.
6394:.
6384:84
6382:.
6361:.
6351:.
6339:.
6335:.
6318:.
6308:29
6306:.
6289:.
6275:.
6271:.
6026:^
5843:^
5745:.
5737:.
5729:.
5719:35
5717:.
5713:.
5690:.
5682:.
5674:.
5662:.
5658:.
5610:^
5581:^
5488:^
5411:^
5384:^
5369:^
5220:^
5205:^
5166:^
5151:^
5052:^
4961:^
4942:^
4873:,
4861:^
4842:^
4813:^
4772:^
4757:^
4728:^
2666:,
2662:,
2539:,
2535:,
2521:,
2515:,
2509:,
2505:,
2499:,
2417:,
2383:.
1906:.
1830:.
1818:,
1761:,
1407:,
1350:.
1325:,
1294:,
1198:,
1010:A
780:,
776:,
770:,
764:,
726:.
228::
163:,
118:Pg
62:Ma
57:,
8046:e
8039:t
8032:v
7975:.
7963::
7946:.
7924::
7916::
7893:.
7869::
7848:.
7834::
7813:.
7791::
7783::
7775::
7754:.
7734::
7726::
7709:.
7689::
7681::
7664:.
7652::
7592:.
7580::
7572::
7566:5
7555:.
7543::
7535::
7494:.
7460::
7452::
7446:4
7424:.
7370:.
7366::
7358::
7334:.
7322::
7297:.
7277::
7260:.
7248::
7240::
7223:.
7195::
7159:.
7138:.
7116::
7095:.
7091::
7085:4
7070:.
7066::
7045:.
7021::
7013::
6979:.
6955::
6947::
6924:.
6918::
6897:.
6875::
6867::
6844:.
6822::
6814::
6808:8
6793:.
6773::
6765::
6744:.
6722::
6716:6
6701:.
6663:.
6642:.
6621:.
6601::
6584:.
6572::
6566:1
6547:.
6526:.
6512::
6506:4
6488:.
6451:.
6402:.
6390::
6369:.
6347::
6341:4
6326:.
6322::
6314::
6297:.
6283::
6258:.
5853:.
5753:.
5733::
5725::
5698:.
5678::
5670::
5664:6
5644:.
4975:.
4877:.
4807:.
3553:â
3510:â
3480:â
3453:â
3435:â
3410:â
3383:â
3365:â
3278:â
3253:â
3222:â
3197:â
3159:â
3124:â
3106:â
3073:â
3042:â
2994:â
2952:â
2891:â
2873:â
2848:â
2801:â
2785:â
2767:â
2729:â
2612:.
2322:'
2316:'
2308:(
2242:(
2230:(
2131:-
1516:(
877:,
693:'
687:'
679:(
602:(
474:(
460:(
307:â
299:â
275:â
265:â
242:â
123:N
113:K
108:J
103:T
98:P
93:C
88:D
83:S
78:O
73:ę
53:-
38:.
20:)
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
