Waeringopteroidea
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Waeringopteridae is a family of
eurypterid Eurypterids, often informally called sea scorpions, are a group of extinct arthropods that form the Order (biology), order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period 467.3 Myr, million yea ...
s, an extinct group of aquatic
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s. The Waeringopteridae is the only family classified as part of the superfamily Waeringopteroidea, which in turn is classified within the infraorder
Diploperculata Diploperculata is an infraorder of eurypterids, an extinct group of aquatic arthropods commonly known as "sea scorpions". The name, derived from Greek διπλόω ("double") and ''operculum'', refers to the distinguishing feature that unites the ...
in the suborder
Eurypterina Eurypterina is one of two suborders of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". Eurypterine eurypterids are sometimes informally known as "swimming eurypterids". They are known from fossil deposi ...
. The earliest known member of the group, '' Orcanopterus'', has been recovered from deposits of
Katian The Katian is the second stage of the Upper Ordovician. It is preceded by the Sandbian and succeeded by the Hirnantian Stage. The Katian began million years ago and lasted for about 7.8 million years until the beginning of the Hirnantian million ...
(
Late Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. The ...
) age and the latest known surviving member, ''
Grossopterus ''Grossopterus'' is a genus of prehistoric eurypterid classified as part of the family Waeringopteridae. The genus contains two species, ''G. inexpectans'' from Gilboa, United States and ''G. overathi'' from Overath, Germany.Dunlop, J. A., Pen ...
'', has been recovered from deposits of Siegenian (
Early Devonian The Early Devonian is the first of three Epoch (geology), epochs comprising the Devonian period, corresponding to the Lower Devonian Series (stratigraphy), series. It lasted from and began with the Lochkovian Stage , which was followed by the P ...
) age. The name Waeringopteridae is derived from the type genus '' Waeringopterus'', which is named in honor of eurypterid researcher Erik N. Kjellesvig-Waering. Waeringopterid fossils are very rare and the family only contains five known species classified in three different genera (''Waeringopterus'', ''Orcanopterus'' and ''Grossopterus''). Four of these species are from the ancient continent of
Laurentia Laurentia or the North American Craton is a large continental craton that forms the ancient geological core of North America. Many times in its past, Laurentia has been a separate continent, as it is now in the form of North America, although ...
, with the only species known from outside of it being a Devonian species recovered from
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. It is thus considered likely that the group originated in Laurentia before spreading to Germany during the Devonian. As the fossil record of the group is extremely poor, little is known of the morphology of the taxa within the group. Although confirmed by phylogenetic analyses as monophyletic, and routinely used within eurypterid taxonomy,Dunlop, J. A., Penney, D. & Jekel, D. 2015. A summary list of fossil spiders and their relatives. In World Spider Catalog. Natural History Museum Bern, online at http://wsc.nmbe.ch, version 16.0 http://www.wsc.nmbe.ch/resources/fossils/Fossils16.0.pdf (PDF). "Waeringopteroidea" and "Waeringopteridae" are as of yet not formally published clades, the names instead deriving from a 2004 thesis by O. Erik Tetlie. Some researchers refer to the group more formally as the "waeringopteroid clade" rather than as the superfamily "Waeringopteroidea", pending a formal publication.


Description

Although waeringopterid fossils remain rare and in most cases fragmentary and very little concrete information on the group can confidently be established due to the poor fossil record, the waeringopterid eurypterids are joined together by a handful of known features shared by all genera within the group. Like all other
chelicerates The subphylum Chelicerata (from New Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. It contains the sea spiders, horseshoe crabs, and arachnids (including harvestmen, scorpions, spiders, solifuges, ticks, and mit ...
, and other arthropods in general, waeringopterid eurypterids possessed segmented bodies and jointed appendages (limbs) covered in a
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
composed of
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
s and
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
. The chelicerate body is divided into two tagmata (sections); the frontal
prosoma The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...
(head) and posterior
opisthosoma The opisthosoma is the posterior part of the body in some arthropods, behind the prosoma (cephalothorax). It is a distinctive feature of the subphylum Chelicerata (arachnids, horseshoe crabs and others). Although it is similar in most respects to a ...
(abdomen). In the waeringopterids, the fifth pair of appendages were spiniferous. The
telson The telson () is the posterior-most division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment of the arthropod body, or an additional division that is not a true segment on accou ...
(the posteriormost segment of the body) was xiphous (long and pointed). The eyes of waeringopterid eurypterids were located close to the marginal rim of the
carapace A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
(the "head" plate). The waeringopterids were almost all small eurypterids. ''Waeringopterus cumberlandicus'' reached 15 cm (5.9 in) in length, while ''Grossopterus overathi'' reached 30 cm (11.8 in). ''Orcanopterus manitoulinensis'' reached 60 cm (23.6 in) in length. Though this is relatively long in comparison to modern arthropods, other members of Diploperculata, such as the
pterygotid Pterygotidae (the name deriving from the type genus ''Pterygotus'', meaning "winged one") is a family of eurypterids, an extinct group of aquatic arthropods. They were members of the superfamily Pterygotioidea. Pterygotids were the largest known ...
s or the carcinosomatids, reached much larger sizes, surpassing two meters.


Classification

The Waeringopteridae is the only family within the superfamily Waeringopteroidea and contains the genera ''
Grossopterus ''Grossopterus'' is a genus of prehistoric eurypterid classified as part of the family Waeringopteridae. The genus contains two species, ''G. inexpectans'' from Gilboa, United States and ''G. overathi'' from Overath, Germany.Dunlop, J. A., Pen ...
'', '' Orcanopterus'' and '' Waeringopterus''. The cladogram below presents the inferred phylogenetic positions of most of the genera included in the three most derived superfamilies of the
Eurypterina Eurypterina is one of two suborders of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". Eurypterine eurypterids are sometimes informally known as "swimming eurypterids". They are known from fossil deposi ...
suborder of eurypterids (
Adelophthalmoidea Adelophthalmidae (the name deriving from the type genus ''Adelophthalmus'', meaning "no obvious eyes") is a family of eurypterids, an extinct group of aquatic arthropods. Adelophthalmidae is the only family classified as part of the superfamily ...
,
Pterygotioidea Pterygotioidea (the name deriving from the type genus ''Pterygotus'', meaning "winged one") is a superfamily of eurypterids, an extinct group of aquatic arthropods. Pterygotioids were the most derived members of the infraorder Diploperculata and ...
and the waeringopteroids), as inferred by O. Erik Tetlie and Markus Poschmann in 2008, based on the results of a 2008 analysis specifically pertaining to the Adelophthalmoidea and a preceding 2004 analysis. "Waeringopteridae" and the superfamily that includes it, "Waeringopteroidea" (named after eurypterid researcher Erik N. Kjellesvig-Waering), are not formally published clades. The names derive from a 2004 thesis by O. Erik Tetlie and are as such not actually technically valid names. Nevertheless, as phylogenetic analyses confirms the grouping as monophyletic and the names supply an easy way to refer to the group, they remain routinely used within studies centered around eurypterid taxonomy. In these studies, the names are usually within quotation marks and/or are noted to not be properly valid names that derive from a thesis. The group is also sometimes referred to as the "waeringopteroid clade" rather than as the "Waeringopteroidea" within studies to differentiate it from the properly described superfamilies, pending the formal publication of Waeringopteroidea.


Distribution and paleobiogeography

Out of the five known waeringopterid species, four are from the fossil deposits in areas that were part of the ancient continent of
Laurentia Laurentia or the North American Craton is a large continental craton that forms the ancient geological core of North America. Many times in its past, Laurentia has been a separate continent, as it is now in the form of North America, although ...
. Only the last known surviving species, '' Grossopterus overathi'', from the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
of
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
, is known from outside of Laurentia and it is thus assumed that the group originated in Laurentia before they spread to modern-day Germany in the Devonian. Further fossil finds are required to properly study the distribution and evolutionary history of the group. The temporal range of the group extends from the
Late Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. The ...
('' Orcanopterus'') to the
Early Devonian The Early Devonian is the first of three Epoch (geology), epochs comprising the Devonian period, corresponding to the Lower Devonian Series (stratigraphy), series. It lasted from and began with the Lochkovian Stage , which was followed by the P ...
(''
Grossopterus ''Grossopterus'' is a genus of prehistoric eurypterid classified as part of the family Waeringopteridae. The genus contains two species, ''G. inexpectans'' from Gilboa, United States and ''G. overathi'' from Overath, Germany.Dunlop, J. A., Pen ...
'').


See also

* List of eurypterid genera


References

{{Taxonbar, from=Q21588877 Diploperculata Silurian first appearances Devonian extinctions Prehistoric arthropod families