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Temporal range: HirnantianPresent, 445–0  Ma
Atlantic horseshoe crab (Limulus polyphemus)
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Clade: Prosomapoda
Order: Xiphosura
Latreille, 1802

Xiphosura ( /ˌzɪfəˈsjʊərə/ ) is an order of arthropods related to arachnids, or, according to one recent study, actual arachnids. [1] 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. They are members of the order Xiphosura, which contains two suborders, Xiphosurida and Synziphosurina.


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 [2] ). Although the name Merostomata is still seen in textbooks, without reference to the Eurypterida, some have urged that this usage should be discouraged. [3] 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. [4] One recent study places Xiphosura within the Arachnida as the sister group of Ricinulei. [1]


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, [5] 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. [6] [7]

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. [7]

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. [8] These are thought to be vestiges of the limbs of an absorbed first opisthosomal segment. [7]

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. [9] The underside of the opisthosoma carries the genital openings and five pairs of flap-like gills. [7]

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. [7]

Internal anatomy

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. [7]

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. [7]

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. [7]


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. [7]

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. [7]


Mesolimulus from the Solnhofen limestone Naturkundemuseum Berlin - Mesolimulus walchi - Solnhofen.jpg
Mesolimulus from the Solnhofen limestone

Xiphosuran classification as of 2018: [10] [11]

Order Xiphosura Latreille, 1802

Taxa removed from Xiphosura

Two groups were originally included in the Xiphosura, but since have been assigned to separate classes:

See also

Related Research Articles

Chelicerata subphylum of arthropods

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.

Arachnid Class of arthropods

Arachnida is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Spiders are the largest order in the class, which also includes scorpions, ticks, mites, harvestmen, and solifuges. In 2019, a molecular phylogenetic study also placed horseshoe crabs in Arachnida.

Ricinulei order of arachnids

The order Ricinulei is a group of arachnids known as hooded tickspiders, though they are not true spiders. Like most arachnids, they are predatory, eating small arthropods. In older works they are sometimes referred to as Podogona.

Eurypterid Order of arthropods (fossil)

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.

Horseshoe crab family of arthropods

Horseshoe crabs are marine and brackish water arthropods of the family Limulidae, suborder Xiphosurida, and order Xiphosura. Their popular name is a misnomer, as they are not true crabs, which are crustaceans.

Trigonotarbida order of arachnids

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.

Chasmataspidid order of arthropods

Chasmataspidids, often referred to informally as chasmataspids, are a group of extinct arthropods that form the order Chasmataspidida. Chasmataspidids are probably related to horseshoe crabs (Xiphosura) and/or sea scorpions (Eurypterida). The first species to be discovered were thought to be unusual fossil horseshoe crabs, while later species were often based on specimens initially misidentified as eurypterids. There is some evidence that chasmataspidids were present during the late Cambrian and the group is known sporadically in the fossil record through to the mid-Devonian. Chasmataspidids are most easily recognised by having an abdomen divided into a short forepart and a longer hindpart comprising nine segments. There is some debate about whether they form a natural group.

<i>Alkenopterus</i> genus of arthropods (fossil)

Alkenopterus is a genus of prehistoric eurypterid classified as part of the family Onychopterellidae. The genus contains two species, A. brevitelson and A. burglahrensis, both from the Devonian of Germany.


Drepanopterus is an extinct genus of eurypterid and the only member of the family Drepanopteridae within the Mycteropoidea superfamily. There are currently three species assigned to the genus. The genus has historically included more species, with nine species associated with the genus Drepanopterus, however five of these have since been proven to be synonyms of pre-existing species, assigned to their own genera, or found to be based on insubstantial fossil data. The holotype of one species proved to be a lithic clast.

<i>Euproops</i> genus of arthropods (fossil)

Euproops is an extinct genus of xiphosuran, related to the modern horseshoe crab. It lived in the Carboniferous.

<i>Unionopterus</i> genus of Eurypterid

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.


The Rhenopteridae are a family of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". The family is the only family currently contained in the superfamily Rhenopteroidea, one of four superfamilies classified as part of the suborder Stylonurina.

Adelophthalmidae A family of eurypterids, an extinct group of aquatic arthropods.

Adelophthalmidae is a family of eurypterids, an extinct group of aquatic arthropods. Adelophthalmidae is the only family classified as part of the superfamily Adelophthalmoidea, which in turn is classified within the infraorder Diploperculata in the suborder Eurypterina.

Eurypterina suborder of eurypterid

Eurypterina is one of two suborders of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". Members of the suborder are collectively and informally known as "eurypterine eurypterids" or "eurypterines". They are known from fossil deposits worldwide, though primarily in North America and Europe.

<i>Strabops</i> genus of extinct arthropod

Strabops is a genus of strabopid, an extinct group of arthropods. Strabops is known from a single specimen from the Late Cambrian of the Potosi Dolomite, Missouri, collected by a former professor, Arthur Thacher. It is classified in the family Strabopidae of the monotypic order Strabopida, a group closely related to the aglaspidids with uncertain affinities. The generic name is composed by the Ancient Greek words στραβός, meaning "squinting", and ὄψῐς, meaning "face".

<i>Borchgrevinkium</i> fossil arthropod genus

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.


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Timeline of eurypterid research

This timeline of eurypterid research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of eurypterids, a group of extinct aquatic arthropods closely related to modern arachnids and horseshoe crabs that lived during the Paleozoic Era.

<i>Dvulikiaspis</i> Extinct genus of chasmataspidid

Dvulikiaspis is a genus of chasmataspidid, a group of extinct aquatic arthropods. Fossils of the single and type species, D. menneri, have been discovered in deposits of the Early Devonian period in the Krasnoyarsk Krai, Siberia, Russia. The name of the genus is composed by the Russian word двуликий (dvulikij), meaning "two-faced", and the Ancient Greek word ἀσπίς (aspis), meaning "shield". The species name honors the discoverer of the holotype of Dvulikiaspis, Vladimir Vasilyevich Menner.


  1. 1 2 Sharma, Prashant P.; Ballesteros, Jesús A. (14 February 2019). "A Critical Appraisal of the Placement of Xiphosura (Chelicerata) with Account of Known Sources of Phylogenetic Error". Systematic Biology. 68 (6): 896–917. doi:10.1093/sysbio/syz011. PMID   30917194.
  2. Garwood, Russell J.; Dunlop, Jason A. (2014). "Three-dimensional reconstruction and the phylogeny of extinct chelicerate orders". PeerJ. 2: e641. doi:10.7717/peerj.641. PMC   4232842 . PMID   25405073.
  3. H. B. Boudreaux (1979). Arthropod Phylogeny with Special Reference to Insects. John Wiley & Sons. pp. 1–320.
  4. Lamsdell, James C. (2012-12-18). "Revised systematics of Palaeozoic 'horseshoe crabs' and the myth of monophyletic Xiphosura". Zoological Journal of the Linnean Society. 167 (1): 1–27. doi: 10.1111/j.1096-3642.2012.00874.x . ISSN   0024-4082.
  5. Crystallographic Texture of the Arthropod Cuticle Using Synchrotron Wide Angle X-ray Diffraction
  6. Botton, M.I. (1984) Diet and food preferences of the adult horseshoe crab Limulus polyphemus in Delaware Bay, New Jersey, USA, Marine Biology, 81, pp. 199-207
  7. 1 2 3 4 5 6 7 8 9 10 Robert D. Barnes (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 590–595. ISBN   978-0-03-056747-6.
  8. R. C. Brusca & G. J. Brusca (2002). Invertebrates. Massachusetts: Sinauer Associates.
  9. Lyall I. Anderson & Paul A. Selden (1997). "Opisthosomal fusion and phylogeny of Palaeozoic Xiphosura". Lethaia . 30 (1): 19–31. doi:10.1111/j.1502-3931.1997.tb00440.x.
  10. Dunlop, J. A., Penney, D. & Jekel, D. 2018. A summary list of fossil spiders and their relatives. In World Spider Catalog. Natural History Museum Bern
  11. Lamsdell, James C. (2016). "Horseshoe crab phylogeny and independent colonizations of fresh water: ecological invasion as a driver for morphological innovation". Palaeontology . 59 (2): 181–194. doi:10.1111/pala.12220.

Further reading