Arthropoda Xiphosura


Chelicerata ├─Merostomata │ ├─Xiphosura │ │ ├─Bunodidae │ │ └─┬─Bellinurina │ │ └─Limulina │ │ ├─Paleolimulidae │ │ └─Limulidae │ └─┬─Chasmataspida │ └─Eurypterida └─Arachnida

   Basal Xiphosura
Links & References


Class Xiphosura Latreille, 1802

(Horseshoe crabs and their extinct relatives)

  Limulus - the horseshoe crab
Limulus polyphemus (Linnaeus)
Limulus photo from Paul D. Bell's Insects Classification - Arthropoda page family Limulidae - "tribe Limulini"


Introduction to the Xiphosura
Order Synziphosurida
family Weinberginidae
family Bunodidae
family Pseudoniscidae
family Kasibelinuridae

Order Xiphosurida
superfamily Bellinuroidea (including Euproopacea)
family Bellinuridae
family Euproopidae
family Elleriidae

superfamily unspecified
family Rolfeiidae

superfamily Limuloidea
family Paleolimulidae
family Moravuridae
family "Valloisellidae"
family Austrolimulidae
family Heterolimulidae
family Limulidae



There are only 3 genera and 5 species of Xiphosura left alive today, but they were much more numerous and diverse during the Palaeozoic era. The surviving horseshoe crabs (Limulus) are 'living fossils', barely changed in some 250 million years (since early Triassic time). Members of this class have a large shield that covers the cephalothorax; the carapace is hinged between the cephalothorax and abdomen. The exoskeleton generally consists of three parts, the large, semicircular cephalothorax, or prosoma, the usually smaller, subtriangular and in earlier forms "trilobite"-like opisthosoma, and the long stout tail-spine or telson (which is actually the end part of the opisthosoma).

The prosoma contains both head and visceral organs. The compound eyes are small (and absent in some early forms), and there are six pairs of legs (in the living Limulus) but no antennae. The second pair of appendages, the pedipalps, resemble walking legs. Respiration is via 5 pairs of book gills, the flaps of which beat in a metachronal rhythm to produce a vigorous current. Recent xiphosurids (Horseshoe crabs) feed on worms and other small invertebrates. They are often used as laboratory animals by physiologists. It has been argued that because of their unique status as prehistoric "living fossils" they deserve special conservation status.


cladogram of Paleozoic Xiphosura

 The Evolution of the Xiphosura during the Paleozoic era, showing representative genera. This cladogram is from Anderson & Selden, reproduced courtesy Lyall Anderson.

Paleomerus and Lemoneites are very early forms that were either Aglaspids or transitional between the Aglaspida and the Xiphosura. The Furongian (Caerfai epoch) marine family Eolimulidae is generally considered a true Xiphosuran, but again more research needs to be done if more is to be known about the early history of this interesting group.


There are only two orders (or sub-orders, depending on your preference) of Xiphosura, the primitive and ancestral Synziphosurina/-ida, and the Lumulina/-ida/Xiphosurida/-ina. The latter group includes modern horseshoe crabs and their immediate ancestors.

The Aglaspida, for a long time considered a very primitive order of Xiphosura, are now thought to be a distinct group, and may actually be closer in fact be closer to the trilobites, or alternatively an ancestral lineage of Merostomata. The Chasmataspida, previously considered Xiphosura, appear to be closer to the Eurypterida, or possibly represent an independent subclass. With modern cladistic analysis even this Linnean arrangement is being discarded.

The following family level Linnean classification presented here is a sort of compromise between the Treatise of Invertebrate Paleontology, the Fossil Record I and II, and the recent work of Lyall Anderson (who presents a cladistic arrangement of the group). In view of the fact that this material is taken hither-thither from other sources it should not be considered an authoritative review of the group.

Order Synziphosurida (Basal Xiphosura)

Silurian to Early Devonian.
Cyamocephalus - Silurian period
Cyamocephalus loganensis Currie 1927
Length about 5 cm - family Pseudoniscidae
Late Llandovery or possibly early Wenlock, Silurian period, Scotland (Euramerica)
image by, and courtesy of, Lyall Anderson.

The Synziohosurida are a small, fairly diverse, paraphyletic / ancestral group of primitive Xiphosura. Rather trilobite-like in appearance. Large prosoma (headshield), simple eyes only. 9 or 10 opisthosomal (abdominal) segments, most or all of which are free (unfused). The segments are not chelate. Mostly brackish or freshwater environments, although some forms were marine (marginal marine?). Usually found in association with eurypterids and ostracoderms.

Family Weinberginidae

WeinberginaFamily Weinberginidae Richter and Richter, 1929
time range: (Pragian to Emsian
habitat: Marine
representative taxa: Weinbergina opitzi Richter and Richter, 1929, Willwerathia laticeps (Størmer, 1936); Legrandella lombardi Eldredge, 1974
status: monophyletic
description: Medium-sized forms with large semicircular smooth prosoma, short 10-segmented Small trilobite-like abdomen, post-abdomen with 3 segments, 5 pairs of walking legs, spines. These late surviving forms seem to be among the most primitive of the Synziphosurida, or in any case they represent a side branch away from the main axis of Xiphosuran evolution. They are also the only Synziphosurines found in a proper marine environment.

Family Bunodidae

family Bunodidae Packard 1886

Small elongate forms, ornamented prosoma, abdomen with broad axis, 9 free segments, post-abdomen with 3 or 4 segments.

Subfamily Limuloidinae

Limuloidesnew ranking; formerly family Limuloididae Størmer, 1952
time range: Wenlock to Ludlow
habitat: Marginal Marine
representative taxa: Limuloides limuloides Woodward, 1865
description: Small forms, ridged and spiny prosoma, post-abdomen with 3 segments. Only one genus.

Subfamily Bunodinae

Bunodesnew ranking; family Bunodidae Packard 1886
time range: Llandovery to Ludlow
habitat: Marginal Marine
representative taxa: Bunodes lunula Eichwald, 1854
description: Small trilobite-like forms, vaulted, radially-lobed prosoma, post-abdomen with 3 to 4 round segments.

Family Pseudoniscidae

Pseudoniscusfamily Pseudoniscidae Packard 1886
time range: Llandovery to Pridoli
habitat: Marginal Marine
representative taxa: Pasternakevia podolica Selden and Drygant, 1987, Cyamocephalus loganensis Currie, 1927, Pseudoniscus aculeatus Nieszkowski, 1859
description: Small forms with large, somewhat flat, smooth prosoma, 9 or 10 free segments with pleural (marginal) spines, no post-abdomen. Pasternakevia is transitional between the Bunodidae and Pseudoniscidae proper.

Family Kasibelinuridae

KasibelinurusFamily Kasibelinuridae Pickett, 1993
time range: Givetian to Famennian
habitat: Marginal Marine
representative taxa: Kasibelinurus amicorum Pickett, 1993
description: The last of the synziphosurnes, and also the most derived (advanced). Live in south-East Gondwanaland (Australia). Presumably ancestral to the Xiphosurida.
status: monospecific

Order Xiphosurida

Devonian to Recent.
Liomesaspis - Carboniferous period
Liomesaspis laevis Raymond 1944
Length about 4 cm
A common coal swamp form - Family Euproopidae
Late Bashkirian ? and early Moscovian to early Gzhelian (Euramerica)
image by, and courtesy of, Lyall Anderson.

The Order Xiphosurida/Limulida includes most Xiphosura, and all post-Devonian forms. These animals were quite common during the Carboniferous, and some forms seem to have been amphibious, although other types (e.g. Paleolimulus) were fully marine. Beginning from small ancestral types they increased in size through the Mesozoic and Cenozoic, and modern horseshoe crabs are giants compared to Paleozoic forms (the horse also has shown a similira tendency to increase in size but reduce in diversity throughout the Tertiary and Quaternary periods), There is the tendency towards fusion of the opisthosomal tergites (free abdominal segments) to form a thoracetron or fused plate. There are several superfamilies and a greater number of families, but only one lineage made it into the post-Paleozoic world. Cladistic analysis so far indicates that the Xiphosurida are a monophyletic taxon.


Suborder Bellinurina

Anderson and Selden consider the Bellinuroidea a suborder (Bellinurina) which contains only two families, the Bellinuridae and the Euproopidae. His Euproopidae would seem to be equivalent to the Treatise's Euproopacea, as it includes the genera Euproops and Liomesaspsis (usually each put in a seperate family of the Euproopacea). These were small animals, with short bodies, only partially fused abdominal segments, and wide rounded horseshoecrab-like headshields, giving them a rather trilobite-like resemblence, apart from the long tail-spine or telson. They frequented the coal swamps and were in all likelihood amphibious, perhaps living part of their life-cycle on land.


time range: Frasnian
habitat: Marginal Marine ? Brackish and Fresh water. Amphibious?
representative taxa:
description: Abdominal axis and pleural area segmented; the rear of the abdominal shield deeply indented. In the Treatise placed in the superfamily Euproopacea. In view of the early date this group may perhaps turn out to be a separate early off-shoot.

Family Euproopidae

EuproopsFamily Euproopidae Eller, 1938 (junior synonym: family Liomesaspidae Raymond 1944
time range: Viséan to Artinskian
habitat: Marginal Marine, Brackish and Fresh water. Amphibious.
representative taxa: Euproops anthrax Prestwich, 1840, Euproops danae Meek and Worthen, 1865, Euproops rotundatus Prestwich, 1840, Liomesaspis laevis Raymond, 1944
description: Small, common forms with well-marked cardiophalmic horseshoe crab "head" area, the cardiac lobe bordered by distinct axial furrows. The abdomen broad and rounded, abdominal shield with annulated (ringed) axis and a boss or knob on the last segment. Like Bellinurids, Euproopids were a common element in the great coal swamps of the Carboniferous tropics.
LiomesaspisTaxanomic note: In the Treatise this family is separated from the Bellinuridae by being placed in the superfamily Euproopacea. Liomesaspis is placed in the family Liomesaspidae, superfamily Euproopacea, and there is also a seperate listing for the family in the Fossil Record II (time range: Bashkirian to Asselian). However Anderson and Selden include Liomesaspis in the family Euproopidae, and Anderson points out that Raymond's initial diagnosis of Liomesaspidae as lacking axial furrows does not apply to the specimens of the genus he has studied. There seems no reason therefore in retained it as a valid family.

time range: Bashkirian to Asselian
habitat: Marginal Marine, Brackish and Fresh water. Amphibious.

Family Bellinuridae

BellinurusFamily Bellinuridae Zittel and Eastman, 1913
time range: Givetian? Serpukhovian to Moscovian
habitat: Marginal Marine, Brackish and Fresh water. Amphibious.
representative taxa: Bellinurus trilobitoides Buckland, 1837.
tentative taxa: Bellinuroopsis rossicus Chernyshev, 1933 (=Neobelinuropsis) may represent a distinct family and superfamily. Protolimulus eriensis Williams) is a small poorly known Devonian form.
description: The most primitive Xiphosurid family, evolving from a Kasibelinurus-like ancestor, representing an ancestral type from which more advanced forms may have developed. The forward abdominal segments are free, but the last two or more rear ones may be fused. This fusing of the abdominal segments is a common and defining tendency among the Xiphosurida.

Suborder Limulina

superfamily Limuloidea Zittel 1885

The Limulina represent the most advanced lineage of the group, descended from early Carboniferous transitional forms like Rolfeia. The cephalothorax is wide and arched, with the cardiophthalmic region poorly defined. .  The abdominal segments are usually fully fused, although the marginal spines are movable. Includes the 5 living species of "horseshoe crabs" (Limulus). These creatures live in a shallow marine environment (although some Paleozoic forms may have been brackish water inhabitants). Jurassic Xiphosurids are extremely similar to those found today.

Anderson and Selden distinguish between the superfamily Paleolimuloidea - defined by the pyramidal cheek node on the carapace - and the superfamily Limuloidea (which Includes all Mesozoic and Cenozoic Xiphosura).

Family Rolfeiidae

RolfeiaRolfeiidae Selden and Siveter, 1987
time range: Tournaisian - Visean
habitat: Marginal Marine
representative taxa: Rolfeia fouldenensis Waterston, 1985
These are paraphyletic basal limulines.

Family Paleolimulidae

PaleolimulusFamily Paleolimulidae Raymond, 1944
time range: Serpukhovian to Asselian (not Hettangian, "Paleolimulus" fuchsbergensis does not belong in the genus)
habitat: Marine, Marginal Marine, Brackish and Fresh water
representative taxa: Paleolimulus avitus Dunbar, 1923
description: Small forms with ophthalmic ridges meeting in front of the cardiac lobe; carapace with pyramidal cheek node; abdominal axis distinctly annulated.

Family Moravuridae

Family Moravuridae Pribyl, 1967

time range: Serpukhovian
habitat: Marginal Marine
representative taxa: Xaniopyramis linseyi Siveter and Selden, 1987

Probably a branch of the Paleolimulidae

Family Undetermined (New family? - "Valloisellidae")

Valloisellatime range: Bashkirian to Moscovian
representative taxa: Valloisella lievensis Racheboeuf, 1992
description: sister taxon to the Limulidae and their relatives. Represent the ancestral type from which modern horseshoe crabs evolved. Basically similar to modern forms.
This, and the following two families are staem limulids, of uncertain monophyly.

Family Austrolimulidae

time range: Ladinian
habitat: Fresh water

Family Heterolimulidae

time range: Ladinian
habitat: Marine

Family Limulidae

Family Limulidae Zittel, 1885
time range: Scythian - Recent
habitat: Marginal Marine, Brackish, and Fresh water
description: Small to large forms; prosoma with ophthalmic ridges not meeting in front of the cardiac lobe; no annulation of abdominal axis; conventional horseshoe crabs.

Subfamily Mesolimulinae Størmer, 1952

In the Treatise this taxon is considered a family.
time range: Scythian to Tithonian (or Cretaceous?)
habitat: Coastal/Marginal Marine, Brackish, and Fresh water
representative taxa: Mesolimulus walchi (Demarest) is known from the Solnhofen late Jurassic. Psammonlimulus gottingensis Lange and Limulitella bronni (Schimper) are early and late Triassic forms respectively.
description: Small to medium-sized forms; genal angles moderately prolonged backwards; axial furrows distinct.

Subfamily Limulinae Zittel, 1885

In the Treatise this taxon is considered a family.
time range: Tertiary to Recent
habitat: Coastal Marine and Brackish water
description: Medium to large forms; hexagonal-shaped abdomen; genal angles strongly prolonged backwards; axial furrows indistinct; "modern" horseshoe crabs.

Tribe Tachypleini Pocock, 1902

new ranking - in the Treatise this taxon is considered a subfamily.
time range: Tertiary to Recent
habitat: Living forms coastal Marine; although the Miocene Tachypleus dechneni of Germany may have inhabited brackish water.
representative taxa: Tachypleus dechneni Zinken (Miocene); Tachypleus gigas (Muller) and Carcinoscorpius rotundatus (Latrieille) both Recent of Asian Pacific.
description: Prosoma slightly vaulted, movable lateral spines of equal length.

Tribe Limulini Zittel, 1885

In the Treatise this taxon is considered a subfamily.
time range: Recent
habitat: Coastal Marine
representative taxa: Limulus polyphemus (Linnaeus)
description: Prosoma vaulted, movable lateral spines decreasing in length backwards.

Links and References

University of California Museum of Paleontology -- Xiphosura - Horseshoe crabs presents the best general introduction to the group on the Web

Lyall I. Anderson's Home Page - excellent material on Paleozoic Xiphosura, and heaps of links

Anderson, LI (1997), The xiphosuran Liomesaspis from the Montceau-les-Mines Konservat Lagerstatte, Massif Central, France. N. Jahrb. Geol. Palaontol., Abh. 204: 415-436.  abstract

Anderson, LI (1998), A new specimen of the Silurian synziphosurine Cyamocephalus. Proc. Geol. Assoc.  110: 211-216. abstract

Anderson, LI & PA Selden (1997), Opisthosomal fusion and phylogeny of Palaeozoic Xiphosura. Lethaia 30: 19-31. abstract

Euproops danae, from the Mazon Creek deposits of Illinois (Moscovian age)  Euproops danae was a small form, length about 2 cm, that is relatively common in the Mazon Creek deposit. It seems to have been freshwater or even semiaquatic, living part of its life on land

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page uploaded 7 May 2002
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last modified ATW060225
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