Eurypterid

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This article is about the order Eurypterida. For the eurypterid family containing
Eurypterus itself, see Eurypteridae.
Eurypterid
Temporal range: Darriwilian-Late Permian, 467.3–251.9 Ma
PreꞒꞒOSDCPTJKPgN
Possible Tremadocian record
Eurypterus remipes 001.jpg
Fossil specimen of Eurypterus remipes housed at the State Museum of Natural History
Karlsruhe in Karlsruhe, Germany.
Scientific classificatione
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Clade: Sclerophorata
Order: †Eurypterida
Burmeister, 1843
Suborders
†Eurypterina Burmeister, 1843
†Stylonurina Diener, 1924
Incertae sedis
Synonyms
Gigantostraca Haeckel, 1866
Cyrtoctenida Størmer & Waterston, 1968
Eurypterids, often informally called sea scorpions, are a group of extinct
arthropods that form the order Eurypterida. The earliest known eurypterids date to
the Darriwilian stage of the Ordovician period 467.3 million years ago. The group
is likely to have appeared first either during the Early Ordovician or Late
Cambrian period. With approximately 250 species, the Eurypterida is the most
diverse Paleozoic chelicerate order. Following their appearance during the
Ordovician, eurypterids became major components of marine faunas during the
Silurian, from which the majority of eurypterid species have been described. The
Silurian genus Eurypterus accounts for more than 90% of all known eurypterid
specimens. Though the group continued to diversify during the subsequent Devonian
period, the eurypterids were heavily affected by the Late Devonian extinction
event. They declined in numbers and diversity until becoming extinct during the
Permian–Triassic extinction event (or sometime shortly before) 251.9 million years
ago.

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

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

Contents
1 Morphology
2 Biology
2.1 Size
2.2 Locomotion
2.3 Respiration
2.4 Ontogeny
2.5 Feeding
2.6 Reproductive biology
3 Evolutionary history
3.1 Origins
3.2 Silurian
3.3 Devonian
3.4 Carboniferous and Permian
4 History of study
5 Classification
5.1 Internal relationships
6 See also
7 References
7.1 Citations
7.2 Bibliography
7.2.1 Websites
8 External links
Morphology

Restoration of Eurypterus with body parts labelled

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

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

The opisthosoma comprised 12 segments and the telson, the posteriormost division of
the body, which in most species took the form of a blade-like shape.[2] In some
lineages, notably the Pterygotioidea, the Hibbertopteridae and the Mycteroptidae,
the telson was flattened and may have been used as a rudder while swimming. Some
genera within the superfamily Carcinosomatoidea, notably Eusarcana, had a telson
similar to that of modern scorpions and may have been capable of using it to inject
venom.[4][5] The coxae of the sixth pair of appendages were overlaid by a plate
that is referred to as the metastoma, originally derived from a complete
exoskeleton segment. The opisthosoma itself can be divided either into a "mesosoma"
(comprising segments 1 to 6) and "metasoma" (comprising segments 7 to 12) or into a
"preabdomen" (generally comprising segments 1 to 7) and "postabdomen" (generally
comprising segments 8 to 12).[2]

The underside of the opisthosoma was covered in structures evolved from modified
opisthosomal appendages. Throughout the opisthosoma, these structures formed plate-
like structures termed Blatfüsse (lit. 'leaf-feet' in German). These created a
branchial chamber (gill tract) between preceding Blatfüsse and the ventral surface
of the opisthosoma itself, which contained the respiratory organs. The second to
sixth opisthosomal segments also contained oval or triangular organs that have been
interpreted as organs that aid in respiration. These organs, termed Kiemenplatten
or "gill tracts", would potentially have aided eurypterids to breath air above
water, while Blatfüssen, similar to organs in modern horseshoe crabs, would cover
the parts that serve for underwater respiration.[2]

The appendages of opisthosomal segments 1 and 2 (the seventh and eighth segments
overall) were fused into a structure termed the genital operculum, occupying most
of the underside of the opisthosomal segment 2. Near the anterior margin of this
structure, the genital appendage (also called the Zipfel 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.[2]

Biology
Size

Size comparison of six of the largest eurypterids: Pterygotus grandidentatus,

Pentecopterus decorahensis, Acutiramus macrophthalmus, A. bohemicus, Carcinosoma
punctatum, and Jaekelopterus rhenaniae
Eurypterids were highly variable in size, depending on factors such as lifestyle,
living environment and taxonomic affinity. Sizes around 100 centimeters (3.3 ft)
are common in most eurypterid groups.[6] The smallest eurypterid, Alkenopterus
burglahrensis, measured just 2.03 centimeters (0.80 in) in length.[7]

The largest eurypterid, and the largest known arthropod ever to have lived, is
Jaekelopterus rhenaniae. A chelicera from the Emsian Klerf Formation of Willwerath,
Germany measured 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 size and body length is
relatively consistent, the specimen of Jaekelopterus 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 of all other known giant arthropods by almost half a meter (1.64 ft) even
if the extended chelicerae are not included.[8] Two other eurypterids have also
been estimated to have reached lengths of 2.5 metres; Erettopterus grandis (closely
related to Jaekelopterus) and Hibbertopterus wittebergensis, but E. grandis is very
fragmentary and the H. wittenbergensis size estimate is based on trackway evidence,
not fossil remains.[9]

The family of Jaekelopterus, the Pterygotidae, is noted for several unusually large
species. Both Acutiramus, whose largest member A. bohemicus measured 2.1 meters
(6.9 ft), and Pterygotus, whose largest species P. grandidentatus measured 1.75
meters (5.7 ft), were gigantic.[8] Several different contributing factors to the
large size of the pterygotids have been suggested, including courtship behaviour,
predation and competition over environmental resources.[10]

Giant eurypterids were not limited to the family Pterygotidae. An isolated 12.7
centimeters (5.0 in) long fossil metastoma of the carcinosomatoid eurypterid
Carcinosoma punctatum indicates the animal would have reached a length of 2.2
meters (7.2 ft) in life, rivalling the pterygotids in size.[11] Another giant was
Pentecopterus decorahensis, a primitive carcinosomatoid, which is estimated to have
reached lengths of 1.7 meters (5.6 ft).[12]

Typical of large eurypterids is a lightweight build. Factors such as locomotion,

energy costs in molting and respiration, as well as the actual physical properties
of the exoskeleton, limits 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.[8] Lightweight adaptations are present in other giant
paleozoic arthropods as well, such as the giant millipede Arthropleura, and are
possibly vital for the evolution of giant size in arthropods.[8][13]

In addition to the lightweight giant eurypterids, some deep-bodied forms in the

family Hibbertopteridae were also very large. A carapace from the Carboniferous of
Scotland referred to the species Hibbertoperus scouleri measures 65 cm (26 in)
wide. As Hibbertopterus was very wide compared to its length, the animal in
question could possibly have measured just short of 2 meters (6.6 ft) in length.
More robust than the pterygotids, this giant Hibbertopterus would possibly have
rivalled the largest pterygotids in weight, if not surpassed them, and as such be
among the heaviest arthropods.[14]

Locomotion

Illustration of subaqueous flight in Eurypterus in which the shape of the paddles

and their motion through water is enough to generate lift. This type of locomotion
was limited to eurypterine (with swimming paddles) eurypterids.

Illustration of Hibbertopterus, a large stylonurine (lacking swimming paddles)

eurypterid
The two eurypterid suborders, Eurypterina and Stylonurina, are distinguished
primarily by the morphology of their final pair of appendages. In the Stylonurina,
this appendage takes the form of a long and slender walking leg, while in the
Eurypterina, the leg is modified and broadened into a swimming paddle.[15] Other
than the swimming paddle, the legs of many eurypterines were far too small to do
much more than allow them to crawl across the sea floor. In contrast, a number of
stylonurines had elongated and powerful legs that might have allowed them to walk
on land (similar to modern crabs).[16]

A fossil trackway was discovered in Carboniferous-aged fossil deposits of Scotland

in 2005 It was attributed to the stylonurine eurypterid Hibbertopterus due to a
matching size (the trackmaker was estimated to have been about 1.6 meters (5.2 ft)
long) and inferred leg anatomy. It is the largest terrestrial trackway—measuring 6
meters (20 ft) long and averaging 95 centimeters (3.12 ft) in width—made by an
arthropod found thus far. It is the first record of land locomotion by a
eurypterid. The trackway provides evidence that some eurypterids could survive in
terrestrial environments, at least for short periods of time, and reveals
information about the stylonurine gait. In Hibbertopterus, as in most eurypterids,
the pairs of appendages are different in size (referred to as a heteropodous limb
condition). These differently sized pairs would have moved in phase, and the short
stride length indicates that Hibbertopterus crawled with an exceptionally slow
speed, at least on land. The large telson was dragged along the ground and left a
large central groove behind the animal. Slopes in the tracks at random intervals
suggest that the motion was jerky.[17] The gait of smaller stylonurines, such as
Parastylonurus, was probably faster and more precise.[18]

The functionality of the eurypterine swimming paddles varied from group to group.
In the Eurypteroidea, the paddles were similar in shape to oars. The condition of
the joints in their appendages ensured their paddles could only be moved in near-
horizontal planes, not upwards or downwards. Some other groups, such as the
Pterygotioidea, would not have possessed this condition and were probably able to
swim faster.[19] Most eurypterines are generally agreed to have utilized a rowing
type of propulsion similar to that of crabs and water beetles. Larger individuals
may have been capable of underwater flying (or subaqueous flight) in which the
motion and shape of the paddles are enough to generate lift, similar to the
swimming of sea turtles and sea lions. This type of movement has a relatively
slower acceleration rate than the rowing type, especially since adults have
proportionally smaller paddles than juveniles. However, since the larger sizes of
adults mean a higher drag coefficient, using this type of propulsion is more
energy-efficient.[20]

The holotype of Palmichnium kosinkiorum, containing the largest eurypterid

footprints known.
Some eurypterines, such as Mixopterus (as inferred from attributed fossil
trackways), were not necessarily good swimmers. It likely kept mostly to the
bottom, using its swimming paddles for occasional bursts of movements vertically,
with the fourth and fifth pairs of appendages positioned backwards to produce minor
movement forwards. While walking, it probably used a gait like that of most modern
insects. The weight of its long abdomen would have been balanced by two heavy and
specialized frontal appendages, and the center of gravity might have been
adjustable by raising and positioning the tail.[21]

Preserved fossilized eurypterid trackways tend to be large and heteropodous and

often have an associated telson drag mark along the mid-line (as with the Scottish
Hibbertopterus track). Such trackways have been discovered on every continent
except for South America. In some places where eurypterid fossil remains are
otherwise rare, such as in South Africa and the rest of the former supercontinent
Gondwana, the discoveries of trackways both predate and outnumber eurypterid body
fossils.[22] Eurypterid trackways have been referred to several ichnogenera, most
notably Palmichnium (defined as a series of four tracks often with an associated
drag mark in the mid-line),[23] wherein the holotype of the ichnospecies P.
kosinkiorum preserves the largest eurypterid footprints known to date with the
found tracks each being about 7.6 centimeters (3.0 in) in diameter.[24] Other
eurypterid ichnogenera include Merostomichnites (though it is likely that many
specimens actually represent trackways of crustaceans) and Arcuites (which
preserves grooves made by the swimming appendages).[23][25][26]

Respiration

The supposed "gill tracts" of eurypterids have been compared to the air-breathing
pseudotracheae present in the posterior legs of modern isopods, such as Oniscus
(pictured).
In eurypterids, the respiratory organs were located on the ventral body wall (the
underside of the opisthosoma). Blatfüsse, evolved from opisthosomal appendages,
covered the underside and created a gill chamber where the "gill tracts" were
located.[2] Depending on the species, the eurypterid gill tract was either
triangular or oval in shape and was possibly raised into a cushion-like state. The
surface of this gill tract bore several spinules (small spines), which resulted in
an enlarged surface area. It was composed of spongy tissue due to many
invaginations in the structure.[27]

Though the Kiemenplatte is referred to as a "gill tract", it may not necessarily

have functioned as actual gills. In other animals, gills are used for oxygen uptake
from water and are outgrowths of the body wall. Despite eurypterids clearly being
primarily aquatic animals that almost certainly evolved underwater (some
eurypterids, such as the pterygotids, would even have been physically unable to
walk on land), it is unlikely the "gill tract" contained functional gills when
comparing the organ to gills in other invertebrates and even fish. Previous
interpretations often identified the eurypterid "gills" as homologous with those of
other groups (hence the terminology), with gas exchange occurring within the spongy
tract and a pattern of branchio-cardiac and dendritic veins (as in related groups)
carrying oxygenated blood into the body. The primary analogy used in previous
studies has been horseshoe crabs, though their gill structure and that of
eurypterids are remarkably different. In horseshoe crabs, the gills are more
complex and composed of many lamellae (plates) which give a larger surface area
used for gas exchange. In addition, the gill tract of eurypterids is proportionally
much too small to support them if it is analogous to the gills of other groups. To
be functional gills, they would have to have been highly efficient and would have
required a highly efficient circulatory system. It is considered unlikely, however,
that these factors would be enough to explain the large discrepancy between gill
tract size and body size.[28]

It has been suggested instead that the "gill tract" was an organ for breathing air,
perhaps actually being a lung, plastron or a pseudotrachea.[29] 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 Blatfüssen.[30] Instead, among
arthropod respiratory organs, the eurypterid gill tracts most closely resemble the
pseudotracheae found in modern isopods. These organs, called pseudotracheae,
because of some resemblance to the tracheae (windpipes) of air-breathing organisms,
are lung-like and present within the pleopods (back legs) of isopods. The structure
of the pseudotracheae has been compared to the spongy structure of the eurypterid
gill tracts. It is possible the two organs functioned in the same way.[31]

Some researchers have suggested that eurypterids may have been adapted to an
amphibious lifestyle, using the full gill tract structure as gills and the
invaginations within it as pseudotrachea. This mode of life may not have been
physiologically possible, however, since water pressure would have forced water
into the invaginations leading to asphyxiation. Furthermore, most eurypterids would
have been aquatic their entire lives. No matter how much time was spent on land,
organs for respiration in underwater environments must have been present. True
gills, expected to have been located within the branchial chamber within the
Blatfüssen, remain unknown in eurypterids.[31]

Ontogeny

Larval (left) and juvenile (right) instars of Strobilopterus (not to scale)

Like all arthropods, eurypterids matured and grew through static developmental
stages referred to as instars. These instars were punctuated by periods during
which eurypterids went through ecdysis (molting of the cuticle) after which they
underwent rapid and immediate growth. Some arthropods, such as insects and many
crustaceans, undergo extreme changes over the course of maturing. Chelicerates,
including eurypterids, are in general considered to be direct developers,
undergoing no extreme changes after hatching (though extra body segments and extra
limbs may be gained over the course of ontogeny in some lineages, such as
xiphosurans and sea spiders). Whether eurypterids were true direct developers (with
hatchlings more or less being identical to adults) or hemianamorphic direct
developers (with extra segments and limbs potentially being added during ontogeny)
has been controversial in the past.[32]

Hemianamorphic direct development has been observed in many arthropod groups, such
as trilobites, megacheirans, basal crustaceans and basal myriapods. True direct
development has on occasion been referred to as a trait unique to arachnids. There
have been few studies on eurypterid ontogeny as there is a general lack of
specimens in the fossil record that can confidently be stated to represent
juveniles.[32] 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.[33]

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

Beartooth Butte Formation in Cottonwood Canyon, Wyoming, composed of multiple
specimens of various developmental stages of eurypterids Jaekelopterus and
Strobilopterus, revealed that eurypterid ontogeny was more or less parallel and
similar to that of extinct and extant xiphosurans, with the largest exception being
that eurypterids hatched with a full set of appendages and opisthosomal segments.
Eurypterids were thus not hemianamorphic direct developers, but true direct
developers like modern arachnids.[34]

The most frequently observed change occurring through ontogeny (except for some
genera, such as Eurypterus, which appear to have been static) is the metastoma
becoming proportionally less wide. This ontogenetic change has been observed in
members of several superfamilies, such as the Eurypteroidea, the Pterygotioidea and
the Moselopteroidea.[35]

Feeding

Pterygotus depicted hunting Birkenia.

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 stereoscopic vision (the ability to perceive depth).
[36] 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 Mixopterus
and Megalograptus). In derived members of the Pterygotioidea, the appendages were
completely without spines, but had specialized claws instead.[37] 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 into their mouth using their chelicerae.[38]

Fossils preserving digestive tracts have been reported from fossils of various
eurypterids, among them Carcinosoma, Acutiramus and Eurypterus. Though a potential
anal opening has been reported from the telson of a specimen of Buffalopterus, it
is more likely that the anus was opened through the thin cuticle between the last
segment before the telson and the telson itself, as in modern horseshoe crabs.[36]

Eurypterid coprolites discovered in deposits of Ordovician age in Ohio containing

fragments of a trilobite and eurypterid Megalograptus ohioensis in association with
full specimens of the same eurypterid species have been suggested to represent
evidence of cannibalism. Similar coprolites referred to the species Lanarkopterus
dolichoschelus from the Ordovician of Ohio contain fragments of jawless fish and
fragments of smaller specimens of Lanarkopterus itself.[36]

Though apex predatory roles would have been limited to the very largest
eurypterids, smaller eurypterids were likely formidable predators in their own
right just like their larger relatives.[6]

Reproductive biology

Type A genital appendage of Adelophthalmus mansfieldi

Type B genital appendage of Kokomopterus longicaudatus
As in many other entirely extinct groups, understanding and researching the
reproduction and sexual dimorphism of eurypterids is difficult, as they are only
known from fossilized shells and carapaces. In some cases, there might not be
enough apparent differences to separate the sexes based on morphology alone.[16]
Sometimes two sexes of the same species have been interpreted as two different
species, as was the case with two species of Drepanopterus (D. bembycoides and D.
lobatus).[39]

The eurypterid prosoma is made up of the first six exoskeleton segments fused
together into a larger structure. The seventh segment (thus the first opisthosomal
segment) is referred to as the metastoma and the eighth segment (distinctly plate-
like) is called the operculum and contains the genital aperature. The underside of
this segment is occupied by the genital operculum, a structure originally evolved
from ancestral seventh and eighth pair of appendages. In its center, as in modern
horseshoe crabs, is a genital appendage. This appendage, an elongated rod with an
internal duct, is found in two distinct morphs, generally referred to as "type A"
and "type B".[16] These genital appendages are often preserved prominently in
fossils and have been the subject of various interpretations of eurypterid
reproduction and sexual dimorphism.[40]

Type A appendages are generally longer than those of type B. In some genera they
are divided into different numbers of sections, such as in Eurypterus where the
type A appendage is divided into three but the type B appendage into only two.[41]
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.[42] The type A appendage is also armed with two curved
spines called furca (lit. 'fork' in Latin). The presence of furca in the type B
appendage is also possible and the structure may represent the unfused tips of the
appendages. Located between the dorsal and ventral surfaces of the Blatfüsse
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 spermathecae (organs for storing sperm), though this function is yet to be
proven conclusively.[43] In arthropods, spermathecae are used to store the
spermatophore received from males. This would imply that the type A appendage is
the female morph and the type B appendage is the male.[16] 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 genitalia).
It is possible that the greater length of the type A appendage means that it was
used as an ovipositor (used to deposit eggs).[44] 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.[2] 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.[45]

The primary function of the long, assumed female, type A appendages was likely to
take up spermatophore from the substrate into the reproductive tract 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 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 furca 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
material.[46]

Type B appendages, assumed male, would have produced, stored and perhaps shaped
spermatophore in a heart-shaped structure on the dorsal surface of the appendage. A
broad genital opening would have allowed large amounts of spermatophore to be
released at once. The long furca associated with type B appendages, perhaps capable
of being lowered like the type A appendage, could have been used to detect whether
a substrate was suitable for spermatophore deposition.[47]

Evolutionary history
Origins

A reconstruction of Pentecopterus, the earliest known eurypterid. The family to

which Pentecopterus belongs, the Megalograptidae, was the first truly successful
eurypterid group.
Until 1882 no eurypterids were known from before the Silurian. Contemporary
discoveries since the 1880s have expanded the knowledge of early eurypterids from
the Ordovician period.[48] The earliest eurypterids known today, the megalograptid
Pentecopterus, date from the Darriwilian stage of the Middle Ordovician, 467.3
million years ago.[49] There are also reports of even earlier fossil eurypterids in
deposits of Late Tremadocian (Early Ordovician) age in Morocco, but these have yet
to be thoroughly studied.[50]

Pentecopterus was a relatively derived eurypterid, part of the megalograptid family

within the carcinosomatoid superfamily. Its derived position suggests that most
eurypterid clades, at least within the eurypterine suborder, had already been
established at this point during the Middle Ordovician.[51] The earliest known
stylonurine eurypterid, Brachyopterus,[6] is also Middle Ordovician in age. The
presence of members of both suborders indicates that primitive stem-eurypterids
would have preceded them, though these are so far unknown in the fossil record. The
presence of several eurypterid clades during the Middle Ordovician suggests that
eurypterids either originated during the Early Ordovician and experienced a rapid
and explosive radiation and diversification soon after the first forms evolved, or
that the group originated much earlier, perhaps during the Cambrian period.[51]

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 Precambrian age,[52] 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 Strabops from the
Cambrian of Missouri,[53] are now classified as aglaspidids or strabopids. The
aglaspidids, once seen as primitive chelicerates, are now seen as a group more
closely related to trilobites.[54]

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 pseudofossils. 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 Laurentia, and demersal (living on the seafloor) and basal animals
from the continents Avalonia and Gondwana.[49] The Laurentian predators, classified
in the family Megalograptidae (compromising the genera Echinognathus, Megalograptus
and Pentecopterus), are likely to represent the first truly successful eurypterid
group, experiencing a small radiation during the Late Ordovician.[55]

Silurian

Reconstruction of Erettopterus, a member of the highly successful Silurian and

Devonian eurypterid family Pterygotidae
Eurypterids were most diverse and abundant between the Middle Silurian and the
Early Devonian, with an absolute peak in diversity during the Pridoli epoch, 423 to
419.2 million years ago, of the very latest Silurian.[15] 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.[48]

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.[56] In most Silurian fossil beds, eurypterine
eurypterids account for 90% of all eurypterids present.[57] Though some were likely
already present by the Late Ordovician (simply missing from the fossil record so
far),[51] a vast majority of eurypterid groups are first recorded in strata of
Silurian age. These include both stylonurine groups such as the Stylonuroidea,
Kokomopteroidea and Mycteropoidea as well as eurypterine groups such as the
Pterygotioidea, Eurypteroidea and Waeringopteroidea.[58]

The most successful eurypterid by far was the Middle to Late Silurian Eurypterus, a
generalist, equally likely to have engaged in predation or scavenging. Thought to
have hunted mainly small and soft-bodied invertebrates, such as worms,[59] species
of the genus (of which the most common is the type species, E. remipes) account for
more than 90% (perhaps as many as 95%) of all known fossil eurypterid specimens.
[57] Despite their vast number, Eurypterus are only known from a relatively short
temporal range, first appearing during the Late Llandovery epoch (around 432
million years ago) and being extinct by the end of the Pridoli epoch.[60]
Eurypterus was also restricted to the continent Euramerica (composed of the
equatorial continents Avalonia, Baltica 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, Eurypterus was limited geographically to
the coastlines and shallow inland seas of Euramerica.[57][61]

During the Late Silurian the pterygotid eurypterids, large and specialized forms
with several new adaptations, such as large and flattened telsons capable of being
used as rudders, and large and specialized chelicerae with enlarged pincers for
handling (and potentially in some cases killing) prey appeared.[3][4] Though the
largest members of the family appeared in the Devonian, large two meter (6.5+ ft)
pterygotids such as Acutiramus were already present during the Late Silurian.[9]
Their ecology ranged from generalized predatory behavior to ambush predation and
some, such as Pterygotus itself, were active apex predators in Late Silurian marine
ecosystems.[62] The pterygotids were also evidently capable of crossing oceans,
becoming one of only two eurypterid groups to achieve a cosmopolitan distribution.
[63]

Devonian

Reconstruction of Adelophthalmus, the only eurypterine (with swimming paddles)

eurypterid to survive the Late Devonian extinction and persist into the subsequent
Carboniferous and Permian periods.
Though the eurypterids continued to be abundant and diversify during the Early
Devonian (for instance leading to the evolution of the pterygotid Jaekelopterus,
the largest of all arthropods), the group was one of many heavily affected by the
Late Devonian extinction. The extinction event, only known to affect marine life
(particularly trilobites, brachiopods and reef-building organisms) effectively
crippled the abundance and diversity previously seen within the eurypterids.[64]

A major decline in diversity had already begun during the Early Devonian and
eurypterids were rare in marine environments by the Late Devonian. During the
Frasnian stage four families went extinct, and the later Famennian saw an
additional five families going extinct.[64] As marine groups were the most
affected, the eurypterids were primarily impacted within the eurypterine suborder.
Only one group of stylonurines (the family Parastylonuridae) went extinct in the
Early Devonian. Only two families of eurypterines survived into the Late Devonian
at all (Adelophthalmidae and Waeringopteridae). The eurypterines experienced their
most major declines in the Early Devonian, during which over 50% of their diversity
was lost in just 10 million years. Stylonurines, on the other hand, persisted
through the period with more or less consistent diversity and abundance but were
affected during the Late Devonian, when many of the older groups were replaced by
new forms in the families Mycteroptidae and Hibbertopteridae.[65]

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 placoderms (armored fish) in both North America and Europe.[66]

Stylonurines of the surviving hibbertopterid and mycteroptid families completely

avoided competition with fish by evolving towards a new and distinct ecological
niche. These families experienced a radiation and diversification through the Late
Devonian and Early Carboniferous, the last ever radiation within the eurypterids,
which gave rise to several new forms capable of "sweep-feeding" (raking through the
substrate in search of prey).[67]

Carboniferous and Permian

Reconstruction of the hibbertopterid Campylocephalus, the last known surviving

eurypterid. Campylocephalus went extinct during the Permian-Triassic extinction
event or sometime shortly before.
Only three eurypterid families—Adelophthalmidae, Hibbertopteridae and Mycteroptidae
—survived the extinction event in its entirety. These were all freshwater animals,
rendering the eurypterids extinct in marine environments.[64] With marine
eurypterid predators gone, sarcopterygian fish, such as the rhizodonts, were the
new apex predators in marine environments.[66] The sole surviving eurypterine
family, Adelophthalmidae, was represented by only a single genus, Adelophthalmus.
The hibbertopterids, mycteroptids and Adelophthalmus survived into the Permian.[68]

Adelophthalmus became the most common of all late Paleozoic eurypterids, existing
in greater number and diversity than surviving stylonurines, and diversified in the
absence of other eurypterines.[69] Out of the 33 species referred to
Adelophthalmus, 23 (69%) are from the Carboniferous alone.[70][71] The genus
reached its peak diversity in the Late Carboniferous. Though Adelophthalmus had
already been relatively widespread and represented around all major landmasses in
the Late Devonian, the amalgamation of Pangaea into a global supercontinent over
the course of the last two periods of the Paleozoic allowed Adelophthalmus to gain
an almost worldwide distribution.[57]

During the Late Carboniferous and Early Permian Adelophthalmus 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 world. With their habitat gone,
Adelophthalmus dwindled in number and had already went extinct by the Leonardian
stage of the Early Permian.[72]

Mycteroptids and hibbertopterids continued to survive for some time, with one genus
of each group known from Permian strata: Hastimima and Campylocephalus
respectively.[73] Hastimima went extinct during the Early Permian,[74] as
Adelophthalmus had, while Campylocephalus persisted longer. A massive incomplete
carapace from Late Permian (Changhsingian stage) deposits in Russia represents the
sole fossil remains of the species C. permianus, which might have reached 1.4
meters (4.6 ft) in length.[9] This giant was the last known surviving eurypterid.
[6] No eurypterids are known from fossil beds higher than the Permian. This
indicates that the last eurypterids died either in the catastrophic extinction
event at its end or at some point shortly before it. This extinction event, the
Permian–Triassic extinction event, is the most devastating mass extinction
recorded, and rendered many other successful Paleozoic groups, such as the
trilobites, extinct.[75]

History of study
See also: Timeline of eurypterid research

Figure of Eurypterus remipes by James E. De Kay (1825).

The first known eurypterid specimen was discovered in the Silurian-aged rocks of
New York, to this day one of the richest eurypterid fossil locations. Samuel L.
Mitchill described the specimen, discovered near Westmoreland in Oneida county in
1818. He erroneously identified the fossil as an example of the fish Silurus,
likely due to the strange, catfish-like appearance of the carapace. Seven years
later, in 1825, James E. DeKay examined the fossil and recognized it as clearly
belonging to an arthropod. He thought the fossil, which he named Eurypterus
remipes, represented a crustacean of the order Branchiopoda, and suggested it might
represent a missing link between the trilobites and more derived branchiopods.[76]
The name Eurypterus derives from Greek eurús (εὐρύς 'broad, wide') and pteron
(πτερόν 'wing').[77]

In 1843, Hermann Burmeister published his view on trilobite taxonomy and how the
group related to other organisms, living and extinct, in the work Die Organisation
der Trilobiten aus ihren lebenden Verwandten entwickelt. He considered the
trilobites to be crustaceans, as previous authors had, and classified them together
with what he assumed to be their closest relatives, Eurypterus and the genus
Cytherina, within a clade he named "Palaeadae". Within Palaeadae, Burmeister
erected three families; the "Trilobitae" (composed of all trilobites), the
"Cytherinidae" (composed only of Cytherina, an animal today seen as an ostracod)
and the Eurypteridae (composed of Eurypterus, then including three species).[78]

The fourth eurypterid genus to be described (following Hibbertopterus in 1836 and

Campylocephalus in 1838, not identified as eurypterids until later), out of those
still seen as taxonomically valid in modern times, was Pterygotus (lit. 'winged
one'), described by Louis Agassiz in 1839.[79] Pterygotus was considerably larger
in size than Eurypterus and when the first fossils were discovered by quarrymen in
Scotland they were referred to as "Seraphims" by the workers. Agassiz first thought
the fossils represented remains of fish, only recognizing their nature as arthropod
remains five years later in 1844.[80]

In 1849, Frederick M'Coy classified Pterygotus together with Eurypterus and

Belinurus (a genus today seen as a xiphosuran) within Burmeister's Eurypteridae.
M'Coy considered the Eurypteridae to be a group of crustaceans within the order
Entomostraca, closely related to horseshoe crabs.[81] A fourth genus, Slimonia,
based on fossil remains previously assigned to a new species of Pterygotus, was
referred to the Eurypteridae in 1856 by David Page.[82]

Evolutionary tree of eurypterids as imagined by John Mason Clarke and Rudolf

Ruedemann in 1912.
Jan Nieszkowski's De Euryptero Remipede (1858) featured an extensive description of
Eurypterus fischeri (now seen as synonymous with another species of Eurypterus, E.
tetragonophthalmus), which, along with the monograph On the Genus Pterygotus by
Thomas Henry Huxley and John William Salter, and an exhaustive description of the
various eurypterids of New York in Volume 3 of the Palaeontology of New York (1859)
by James Hall, contributed massively to the understanding of eurypterid diversity
and biology. These publications were the first to fully describe the whole anatomy
of eurypterids, recognizing the full number of prosomal appendages and the number
of preabdominal and postabdominal segments. Both Nieszkowski and Hall recognized
that the eurypterids were closely related to modern chelicerates, such as horseshoe
crabs.[83]

In 1865, Henry Woodward described the genus Stylonurus (named and figured, but not
thoroughly described, by David Page in 1856) and raised the rank of the
Eurypteridae to that of order, effectively creating the Eurypterida as the
taxonomic unit it is seen as today.[84] In the work Anatomy and Relations of the
Eurypterida (1893), Malcolm Laurie added considerably to the knowledge and
discussion of eurypterid anatomy and relations. He focused on how the eurypterids
related to each other and to trilobites, crustaceans, scorpions, other arachnids
and horseshoe crabs. The description of Eurypterus fischeri by Gerhard Holm in 1896
was so elaborate that the species became one of the most completely known of all
extinct animals, so much so that the knowledge of E. fischeri was comparable with
the knowledge of its modern relatives (such as the Atlantic horseshoe crab). The
description also helped solidify the close relationship between the eurypterids and
other chelicerates by showcasing numerous homologies between the two groups.[85]

In 1912, John Mason Clarke and Rudolf Ruedemann published The Eurypterida of New
York 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 Eurypterus, Anthraconectes, Stylonurus, Eusarcus,
Dolichopterus, Onychopterus and Drepanopterus) and Pterygotidae (distinguished by
faceted eyes and including Pterygotus, Erettopterus, Slimonia and Hughmilleria).
Both families were considered to be descended from a common ancestor, Strabops.[86]
In line with earlier authors, Clarke and Ruedemann also supported a close
relationship between the eurypterids and the horseshoe crabs (united under the
class Merostomata) but also discussed alternative hypotheses such as a closer
relation to arachnids.[87]

Classification

Eurypterids have historically been seen as closely related to xiphosurans such as

the Atlantic horseshoe crab (top), united in the class Merostomata. Recent studies
are in favor of a closer relationship to arachnids such as Heterophrynus (bottom),
united under the clade Sclerophorata.
Historically, a close relationship between eurypterids and xiphosurans (such as the
modern Atlantic horseshoe crab) has been assumed by most researchers. Several
homologies encourage this view, such as correlating segments of the appendages and
the prosoma. Additionally, the presence of plate-like appendages bearing the "gill
tracts" on appendages of the opisthosoma (the Blatfüssen) was cited early as an
important homology. In the last few decades of the nineteenth century, further
homologies were established, such as the similar structures of the compound eyes of
Pterygotus and horseshoe crabs (seen as especially decisive as the eye of the
horseshoe crab was seen as possessing an almost unique structure) and similarities
in the ontogeny within both groups.[88] These ontogenetical similarities were seen
as most apparent when studying the nepionic stages (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.[89]

Due to these similarities, the xiphosurans and eurypterids have often been united
under a single class or subclass 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 the horseshoe crabs and the arachnids.[89] Some authors, such as
John Sterling Kingsley in 1894, classified the Merostomata as a sister group to the
Arachnida under the class "Acerata" within a subphylum "Branchiata". Others, such
as Ray Lankester in 1909, went further and classified the Merostomata as a subclass
within the Arachnida, raised to the rank of class.[90]

In 1866, Ernst Haeckel classified the Merostomata (containing virtually only the
Eurypterida) and Xiphosura within a group he named Gigantostraca within the
crustaceans. Though Haeckel did not designate any taxonomic rank for this clade, it
was interpreted as equivalent to the rank of subclass, such as the Malacostraca and
Entomostraca, by later researchers such as John Sterling Kinsgsley.[91] In
subsequent research, Gigantostraca has been treated as synonymous with Merostomata
(rarely) and Eurypterida itself (more commonly).[92][93]

Restoration of the primitive Cambrian arachnomorph Fuxianhuia, one of the most

basal members of the lineage that would eventually lead to both the chelicerates
and the trilobites.
A phylogenetic analysis (the results presented in a cladogram below) conducted by
James Lamsdell in 2013 on the relationships within the Xiphosura and the relations
to other closely related groups (including the eurypterids, which were represented
in the analysis by genera Eurypterus, Parastylonurus, Rhenopterus and
Stoermeropterus) concluded that the Xiphosura, as presently understood, was
paraphyletic (a group sharing a last common ancestor but not including all
descendants of this ancestor) and thus not a valid phylogenetic group.[94]
Eurypterids were recovered as closely related to arachnids instead of xiphosurans,
forming the group Sclerophorata within the clade Dekatriata (composed of
sclerophorates and chasmataspidids). 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 Prosomapoda, a group including
the Xiphosurida (the only monophyletic xiphosuran group) and other stem-genera.[95]

Arachnomorpha
 
†Fuxianhuia

 
 
†Antennulata
 
†Emeraldella

 
 
†Trilobitomorpha

 
 
†Sidneyia

 
 
 
†Megacheira
 
†Yohoia

 
 
 
†Alalcomenaeus

 
 
†Leanchoilia

 
 
 
Chelicerata
Pycnogonida
 
†Palaeoisopus

 
 
 
Pycnogonum

 
 
†Haliestes

 
 
 
Euchelicerata
 
†Offacolus

 
Prosomapoda
 
†Weinbergina

 
 
 
 
†Venustulus

 
 
†Camanchia

 
 
 
 
†Legrandella

 
 
Xiphosura
 
†Kasibelinurus

 
 
 
†Willwerathia

 
Xiphosurida
 
†Lunataspis

 
 
 
†Belinurina

 
 
Limulina

 
 
 
 
 
Planaterga
 
 
†Pseudoniscus

 
 
†Cyamocephalus

 
 
 
†Pasternakevia

 
 
 
†Bunodes

 
 
†Limuloides

 
 
 
†Bembicosoma

 
Dekatriata
 
†Chasmataspidida
 
Sclerophorata
 
Arachnida

 
 
†Eurypterida

 
 
 
 
 
 
 
 
 
 
 
 
 
Internal relationships

The most important taxonomic character in eurypterids is the morphology of the

sixth pair of appendages. In most eurypterids of the suborder Eurypterina, such as
Bassipterus (top), this leg is modified into a swimming paddle. In eurypterids of
the suborder Stylonurina, such as Parastylonurus (bottom), it is not.
The internal classification of eurypterids within the Eurypterida is based mainly
on eleven established characters. These have been used throughout the history of
eurypterid research to establish clades and genera. These characters include: the
shape of the prosoma, the shape of the metastoma, the shape and position of the
eyes, the types of prosomal appendages, the types of swimming leg paddles, the
structure of the doublure (the fringe of the dorsal exoskeleton), the structure of
the opithosoma, the structure of the genital appendages, the shape of the telson
and the type of ornamentation present. It is worth noting that not all of these
characters are of equal taxonomic importance.[96] They are not applicable to all
eurypterids either; stylonurine eurypterids lack swimming leg paddles entirely.[15]
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.[97] Most superfamilies and families are defined based on the
morphology of the appendages.[98]

The most important character used in eurypterid taxonomy is the type of prosomal
appendages as this character is used to define entire suborders. General leg
anatomy can also be used to define superfamilies and families. Historically, the
chelicerae were considered the most important appendages from a taxonomical
standpoint since they only occurred in two general types: a eurypterid type with
small and toothless pincers and a pterygotid type with large pincers and teeth.
This distinction has historically been used to divide the Eurypterida into the two
suborders Eurypterina (small chelicerae) and "Pterygotina" (large and powerful
chelicerae).[99] 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.[100]
Modern research favors a classification into suborders Eurypterina and Stylonurina
instead, supported by phylogenetic analyses.[101][35] In particular, pterygotid
eurypterids share a number of homologies with derived eurypterine eurypterids such
as the adelophthalmids, and are thus best classified as derived members of the same
suborder.[102] 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.
[15] 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.[57]

The cladogram presented below, covering all currently recognized eurypterid

families, follows a 2007 study by O. Erik Tetlie.[103] The stylonurine suborder
follows a 2010 study by James Lamsdell, Simon J. Braddy and Tetlie.[104] The
superfamily "Megalograptoidea", recognized by Tetlie in 2007 and then placed
between the Onychopterelloidea 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.[105]

Eurypterida
Stylonurina
 
Rhenopteridae Brachyopterus restoration.png

 
 
Stylonuroidea
 
Parastylonuridae

 
 
Stylonuridae Stylonurus BW white background.png

 
 
 
Kokomopteroidea
 
Hardieopteridae

 
 
Kokomopteridae

 
 
Mycteropoidea
 
Drepanopteridae

 
 
 
Hibbertopteridae Hibbertopterus scouleri.jpg
 
 
Mycteroptidae Megarachne BW.jpg

 
 
 
 
 
 
Eurypterina
 
Moselopteridae Stoermeropterus.png

 
 
 
Onychopterellidae

 
 
Eurypteroidea
 
Dolichopteridae

 
 
 
Eurypteridae Eurypterus Paleoart (no background).png

 
 
Strobilopteridae

 
 
 
Diploperculata
Carcinosomatoidea
 
Megalograptidae Megalograptus color reconstruction.png

 
 
 
Carcinosomatidae Carcinosoma newlini.png

 
 
Mixopteridae Mixopterus BW.jpg

 
 
 
 
 
Waeringopteridae
 
 
 
Adelophthalmidae Adelophthalmus irinae.png

 
Pterygotioidea
 
Hughmilleriidae

 
 
 
Pterygotidae Pterygotus anglicus reconstruction.jpg

 
 
Slimonidae Slimonia acuminata reconstruction.jpg