|Atlantic horseshoe crab (Limulus polyphemus)|
|Order:|| Xiphosura |
Xiphosura ( // ) is an order of arthropods related to arachnids. They are sometimes called horseshoe crabs (a name applied more specifically to the only extant family, Limulidae). They first appeared in the Hirnantian (Late Ordovician). Currently, there are only four living species. Xiphosura contains one suborder, Xiphosurida, and several stem-genera.
The group has hardly changed in hundreds of millions of years; the modern horseshoe crabs look almost identical to prehistoric genera such as the Jurassic Mesolimulus , and are considered to be living fossils. The most notable difference between ancient and modern forms is that the abdominal segments in present species are fused into a single unit in adults.
Xiphosura were traditionally placed in the class Merostomata, although this term was intended to encompass also the eurypterids, whence it denoted what is now known to be an unnatural (paraphyletic) group (although this is a grouping recovered in some recent cladistic analyses). Although the name Merostomata is still seen in textbooks, without reference to the Eurypterida, some have urged that this usage should be discouraged. The Merostomata label originally did not include Eurypterida, although they were added in as a better understanding of the extinct group evolved. Now Eurypterida is classified within Sclerophorata together with the arachnids, and therefore, Merostomata is now a synonym of Xiphosura. One recent study places Xiphosura within the Arachnida as the sister group of Ricinulei.
Modern xiphosurans reach up to 60 cm (24 in) in adult length, but the Paleozoic species were often far smaller, some as small as 1 to 3 cm (0.39 to 1.18 in) long.
Their bodies are covered with a tough cuticle, but do not contain any crystalline biominerals,and are divided into an anterior prosoma and a posterior opisthosoma, or abdomen. The upper surface of the prosoma is covered by a semicircular carapace, while the underside bears five pairs of walking legs and a pair of pincer-like chelicerae. The mouth is located on underside of the center of the prosoma, between the bases of the walking legs, and lies behind a lip-like structure called the labrum.
Xiphosurans have up to four eyes, located in the carapace. Two compound eyes are on the side of the prosoma, with one or two median ocelli towards the front. The compound eyes are simpler in structure than those of other arthropods, with the individual ommatidia not being arranged in a compact pattern. They can probably detect movement, but are unlikely to be able to form a true image. In front of the ocelli is an additional organ that probably functions as a chemoreceptor.
The first four pairs of legs end in pincers, and have a series of spines, called the gnathobase, on the inner surface. The spines are used to masticate the food, tearing it up before passing it to the mouth. The fifth and final pair of legs, however, has no pincers or spines, instead having structures for cleaning the gills and pushing mud out of the way while burrowing. Behind the walking legs is a sixth set of appendages, the chilaria, which are greatly reduced in size and covered in hairs and spines.These are thought to be vestiges of the limbs of an absorbed first opisthosomal segment.
The opisthosoma is divided into a forward mesosoma, with flattened appendages, and a metasoma at the rear, which has no appendages. In modern forms, the whole of the opisthosoma is fused into a single unsegmented structure.The underside of the opisthosoma carries the genital openings and five pairs of flap-like gills.
The opisthosoma terminates in a long caudal spine, commonly referred to as a telson (though this same term is also used for a different structure in crustaceans). The spine is highly mobile, and is used to push the animal upright if it is accidentally turned over.
The mouth opens into a sclerotised oesophagus, which leads to a crop and gizzard. After grinding up its food in the gizzard, the animal regurgitates any inedible portions, and passes the remainder to the true stomach. The stomach secretes digestive enzymes, and is attached to an intestine and two large caeca that extend through much of the body, and absorb the nutrients from the food. The intestine terminates in a sclerotised rectum, which opens just in front of the base of the caudal spine.
Xiphosurans have well-developed circulatory systems, with numerous arteries that send blood from the long tubular heart to the body tissues, and then to two longitudinal sinuses next to the gills. After being oxygenated, the blood flows into the body cavity, and back to the heart. The blood contains haemocyanin, a blue copper-based pigment performing the same function as haemoglobin in vertebrates, and also has blood cells that aid in clotting.
The excretory system consists of two pairs of coxal glands connected to a bladder that opens near the base of the last pair of walking legs. The brain is relatively large, and, as in many arthropods, surrounds the oesophagus. In both sexes, the single gonad lies next to the intestine and opens on the underside of the opisthosoma.
Xiphosurans move to shallow water to mate. The male climbs onto the back of the female, gripping her with his first pair of walking legs. The female digs out a depression in the sand, and lays from 200 to 300 eggs, which the male covers with sperm. The pair then separates, and the female buries the eggs.
The egg is about 2–3 mm (0.08–0.12 in) across, and hatches into a larva that superficially resembles a trilobite. Indeed, it is often referred to as the 'trilobite larva'. Through a series of successive moults, the larva develops additional gills, increases the length of its caudal spine, and gradually assumes the adult form. Modern xiphosurans reach sexual maturity after about three years of growth.
Xiphosuran classification as of 2018 [update] :
Order Xiphosura Latreille, 1802
Two groups were originally included in the Xiphosura, but since have been assigned to separate classes:
The subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda. It contains the sea spiders, arachnids, and several extinct lineages, such as the eurypterids and chasmataspidids.
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 251.9 million years ago.
The order Trigonotarbida is a group of extinct arachnids whose fossil record extends from the late Silurian to the early Permian. These animals are known from several localities in Europe and North America, as well as a single record from Argentina. Trigonotarbids can be envisaged as spider-like arachnids, but without silk-producing spinnerets. They ranged in size from a few millimetres to a few centimetres in body length and had segmented abdomens, with the tergites across the backs of the animals' abdomens, which were characteristically divided into three or five separate plates. Probably living as predators on other arthropods, some later trigonotarbid species were quite heavily armoured and protected themselves with spines and tubercles. About seventy species are currently known, with most fossils originating from the Carboniferous coal measures. In July 2014 scientists used computer graphics to re-create a possible walking gait for the animal. However, a subsequent biomechanical analysis proved the proposed coordination pattern as largely non-physiological.
Chasmataspidids, sometime referred to as chasmataspids, are a group of extinct chelicerate arthropods that form the order Chasmataspidida. Chasmataspidids are probably related to horseshoe crabs (Xiphosura) and/or sea scorpions (Eurypterida), with more recent studies suggest that they form a clade (Dekatriata) with Eurypterida and Arachnida. Chasmataspidids are known sporadically in the fossil record through to the mid-Devonian, with possible evidence suggest that they also present during the late Cambrian. Chasmataspidids are most easily recognised by having an abdomen divided into a short forepart (preabdomen) and a longer hindpart (postabdomen) each comprising 4 and 9 segments respectively. There is some debate about whether they form a natural group.
Willwerathia is a genus of synziphosurine, a paraphyletic group of horseshoe crab-like fossil chelicerate arthropods. Willwerathia known only by one species, Willwerathia laticeps, discovered in deposits of the Devonian period from the Rhenish Slate Mountains of Germany.
Unionopterus is a genus of eurypterid, an extinct group of aquatic arthropods commonly known as "sea scorpions". Fossils have been registered from the Early Carboniferous period. The genus contains only one species, U. anastasiae, recovered from deposits of Tournaisian to Viséan stages in Kazakhstan. Known from one single specimen which was described in a publication of Russian language with poor illustrations, Unionopterus' affinities are extremely poorly known.
Bembicosoma is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Bembicosoma was regarded as part of the clade Planaterga. Fossils of the single and type species, B. pomphicus, have been discovered in deposits of the Silurian period in the Pentland Hills, Scotland. Bembicosoma had been tentatively assigned as an eurypterid before its synziphosurine affinities revealed.
Bunodes is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Bunodes was regarded as part of the clade planaterga. Fossils of the single and type species, B. lunula, have been discovered in deposits of the Silurian period in Ludlow, England. Bunodes is the type genus of the family Bunodidae, the other genera of the same family being Limuloides.
Cyamocephalus is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Cyamocephalus was regarded as part of the clade Planaterga. Fossils of the single and type species, C. loganensis, have been discovered in deposits of the Silurian period in Lesmahagow, Scotland. Cyamocephalus is one of the two members of the family Pseudoniscidae, the other being Pseudoniscus. Cyamocephalus differ from Pseudoniscus by the fused tergites of 6th and 7th opisthosomal segments.
Legrandella is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Legrandella was regarded as part of the clade Prosomapoda. Fossils of the single and type species, L. lombardii, have been discovered in deposits of the Devonian period in Cochabamba, Bolivia.
Limuloides is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Limuloides was regarded as part of the clade Planaterga. Fossils of the genus have been discovered in deposits of the Silurian period in the United Kingdom and potentially in the United States. Limuloides is one of the two genera of the family Bunodidae, the other being the type genus Bunodes. Limuloides characterized by a carapace with radiated ridges and serrated lateral regions, as well as an opisthosoma with rows of nodes. Limuloides was once though to have lateral compound eyes on its carapace, but later investigation did not find any evidence of it.
Pasternakevia is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Pasternakevia was regarded as part of the clade Planaterga. Fossils of the single and type species, P. podolica, have been discovered in deposits of the Silurian period in Podolia, Ukraine.
Pseudoniscus is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Pseudoniscus was regarded as part of the clade Planaterga. Fossils of the genus have been discovered in deposits of the Silurian period in the United Kingdom, the United States and Estonia. Pseudoniscus is one of the two members of the family Pseudoniscidae, the other being Cyamocephalus.
Weinbergina is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Fossils of the single and type species, W. opitzi, have been discovered in deposits of the Devonian period in the Hunsrück Slate, Germany.
Synziphosurina is a paraphyletic group of chelicerate arthropods previously thought to be basal horseshoe crabs (Xiphosura). It was later identified as a grade compose of various basal euchelicerates, eventually excluded form the monophyletic Xiphosura sensu stricto and only regarded as horseshoe crabs under a broader sense. Synziphosurines survived at least since early Ordovician to early Carboniferous in ages, with most species are known from the in-between Silurian strata.
Borchgrevinkium is an extinct genus of chelicerate arthropod. A fossil of the single and type species, B. taimyrensis, has been discovered in deposits of the Early Devonian period in the Krasnoyarsk Krai, Siberia, Russia. The name of the genus honors Carsten Borchgrevink, an Anglo-Norwegian explorer who participated in many expeditions to Antarctica. Borchgrevinkium represents a poorly known genus whose affinities are uncertain.
The metastoma is a ventral single plate located in the opisthosoma of non-arachnid dekatriatan chelicerates such as eurypterids, chasmataspidids and the genus Houia. The metastoma located between the base of 6th prosomal appendage pair and may had functioned as part of the animal's feeding structures. It most likely represented a fused appendage pair originated from somite 7, thus homologous to the chilaria of horseshoe crab and 4th walking leg pair of sea spider.
Dekatriata is a clade of planatergan chelicerates including the groups Arachnida, Chasmataspidida, Eurypterida and additionally two stem-genera Winneshiekia and Houia. Dekatriata is defined by an opisthosoma with 13 segments as groundplan and fused, plate-like appendages on the first opisthosomal segment.
Camanchia is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Camanchia was regarded as part of the clade Prosomapoda. Fossils of the single and type species, C. grovensis, have been discovered in deposits of the Silurian period in Iowa, in the United States. Alongside Venustulus, Camanchia is one of the only Silurian synziphosurine with fossil showing evidence of appendages.
Anderella is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Anderella was regarded as part of the clade Prosomapoda. Fossils of the single and type species, A. parva, have been discovered in deposits of the Carboniferous period in Montana, in the United States. Anderella is the first and so far the only Carboniferous synziphosurine being described, making it the youngest member of synziphosurines. Anderella is also one of the few synziphosurine genera with fossil showing evidence of appendages, but the details are obscure due to their poor preservation.
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