Horseshoe crab

Horseshoe crabs are marine and brackish water arthropods of the

family Limulidae, suborder Xiphosurida, and order Xiphosura.[3] Limulidae
Their popular name is a misnomer, as they are not true crabs, nor Temporal range: 244–0 Ma [1]
even crustaceans, as crabs are, but a different order of arthropod. PreꞒ Ꞓ O S D C P T J K PgN

Horseshoe crabs live primarily in and around shallow coastal waters

on soft sandy or muddy bottoms. They tend to spawn in the
intertidal zone at spring high tides.[4] They are commonly eaten in
Asia, and used as fishing bait, in fertilizer and in science (especially
Limulus amebocyte lysate). In recent years, population declines
have occurred as a consequence of coastal habitat destruction and
overharvesting.[3] Tetrodotoxin may be present in one horseshoe
crab species, Carcinoscorpius rotundicauda.[5]
Limulus polyphemus
Given their origin 450 million years ago, horseshoe crabs are
considered living fossils.[6] A 2019 molecular analysis places them Scientific classification
as the sister group of Ricinulei within Arachnida.[7] Kingdom: Animalia
Phylum: Arthropoda

Contents Subphylum: Chelicerata

Order: Xiphosura
Taxonomy
Anatomy and behavior Suborder: Xiphosurida

Breeding Family: Limulidae

Leach, 1819 [2]
Threats
Harvest for blood Genera
Fishery
Shoreline development Carcinoscorpius
References Limulus
Further reading †Mesolimulus
External links Tachypleus

Taxonomy
Horseshoe crabs resemble crustaceans but belong to a separate subphylum of the arthropods, Chelicerata.[8]
Horseshoe crabs are closely related to the extinct eurypterids (sea scorpions), which include some of the largest
arthropods to have ever existed, and the two may be sister groups.[8][9] Other studies have placed eurypterids
closer to the arachnids in a group called Merostomata.[10] The enigmatic Chasmataspidids are also thought to
be closely related to the horseshoe crabs.[11] The earliest horseshoe crab fossils are found in strata from the late
Ordovician period, roughly 450 million years ago.
The Limulidae are the only recent family of the order Xiphosura, and contains all four living species of
horseshoe crabs:[2][3]

Carcinoscorpius rotundicauda, the mangrove horseshoe crab, found in South and Southeast
Asia
Limulus polyphemus, the Atlantic or American horseshoe crab, found along the American
Atlantic coast and in the Gulf of Mexico
Tachypleus gigas, the Indo-Pacific, Indonesian, Indian or southern horseshoe crab, found in
South and Southeast Asia
Tachypleus tridentatus, the Chinese, Japanese or tri-spine horseshoe crab, found in Southeast
and East Asia

Anatomy and behavior

The entire body of the horseshoe crab is protected by a hard carapace.
It has two compound lateral eyes, each composed of about 1,000
ommatidia, plus a pair of median eyes that are able to detect both
visible light and ultraviolet light, a single endoparietal eye, and a pair
of rudimentary lateral eyes on the top. The latter become functional
just before the embryo hatches. Also, a pair of ventral eyes is located
near the mouth, as well as a cluster of photoreceptors on the telson.
Having relatively poor eyesight, the animals have the largest rods and
cones of any known animal, about 100 times the size of
Underside of two horseshoe crabs humans',[12][13] and their eyes are a million times more sensitive to
showing the legs and book gills light at night than during the day.[14] They use their chelicerae—a pair
of small appendages—for moving food into the mouth. The next five
pairs of appendages, the first of which are the pedipalps, are used for
locomotion (ambulatory legs). The mouth is located in the center of the legs, whose bases are referred to as
gnathobases. and have the same function as jaws and help grind up food.[15] In extant species their
appendages are uniramous, but the fossil species Dibasterium had four pairs of branched walking legs.[16] The
pedipalps on a male change shape on their terminal molt, becoming boxing glove-like claspers that are used for
grasping the female during mating. The last pair of legs for both male and female are the main legs used for
pushing when walking on the ocean floor. The remaining leg pairs have a weak claw at the tip.[17] Lost legs or
the telson (tail) may slowly regenerate, and cracks in the body shell can heal.[18]

Behind its legs, the horseshoe crab has book gills, which exchange External video
respiratory gases, and are also occasionally used for swimming.[19]
As in other arthropods, a true endoskeleton is absent, but the body
does have an endoskeletal structure made up of cartilaginous plates
that support the book gills. They are more often found on the ocean
floor searching for worms and molluscs, which are their main food.
They may also feed on crustaceans and even small fish.

Females are about 20–30% larger than males.[20] The smallest

species is C. rotundicauda and the largest is T. tridentatus.[21] On Rendezvous with a Horseshoe
average, males of C. rotundicauda are about 30 centimetres (12 Crab (https://www.youtube.com/wat
inches) long, including a tail (telson) that is about 15 cm (6 in), and ch?v=6gydJh6rP50), August 2011,
their carapace (prosoma) is about 15 cm (6 in) wide.[22] Some
4:34, NewsWorks
southern populations (in the Yucatán Peninsula) of L. polyphemus
are somewhat smaller, but otherwise this species is larger.[20] In the The Horseshoe Crab Spawn (htt
largest species, T. tridentatus, females can reach as much as 79.5 cm ps://www.youtube.com/watch?v=wA
(311 ⁄4 in) long, including their tail, and up to 4 kg (9 lb) in -NoH_G330), June 2010, 5:08,
weight.[23] This is only about 10–20 cm (4–8 in) longer than the HostOurCoast.com
largest females of L. polyphemus and T. gigas, but roughly twice the Horseshoe Crabs Mate in
weight.[24][25] The juveniles grow about 33% larger with every molt Massive Beach "Orgy" (https://www.
until reaching adult size.[26] Atlantic horseshoe crabs molt in late youtube.com/watch?v=-55qGCHx1l
July.
E&list=TLzuAKoEqoVX1TChQbCu_
hh27r4Lw-bGmZ), June 2014, 3:29,
National Geographic

Horseshoe crabs normally swim upside
down, inclined at about 30° to the
horizontal and moving at about 10-15
cm/s.[27][28][29]
Horseshoe crabs have two primary
compound eyes and seven
secondary simple eyes. Two of the
secondary eyes are on the
underside.[30][31]

Painting by Heinrich Harder, c. 1916

Breeding
During the breeding season, horseshoe crabs migrate to shallow coastal waters. A male selects a female and
clings to her back. Often, several males surround the female and all fertilize together, which makes it easy to
spot and count females as they are the large center carapace surrounded by 3-5 smaller ones. The female digs a
hole in the sand and lays her eggs while the male(s) fertilize them. The female can lay between 60,000 and
120,000 eggs in batches of a few thousand at a time. In L. polyphemus, the eggs take about two weeks to
hatch; shore birds eat many of them before they hatch. The larvae molt six times during the first year and
annually after the first 3 or 4 years.[32][33]
Natural breeding of horseshoe crabs in captivity has proven to be
difficult. Some evidence indicates that mating takes place only in the
presence of the sand or mud in which the horseshoe crab's eggs were
hatched. It is not known with certainty what is in the sand that the
crabs can sense or how they sense it.[34] Artificial insemination and
induced spawning have been done on a relatively large scale in
captivity, and eggs and juveniles collected from the wild are often
raised to adulthood in captivity.[35][36]
Horseshoe crabs mating
Threats

Harvest for blood

Horseshoe crabs use hemocyanin to carry oxygen through their blood.

Because of the copper present in hemocyanin, their blood is blue.[37]
Their blood contains amebocytes, which play a similar role to the
white blood cells of vertebrates in defending the organism against
pathogens. Amebocytes from the blood of L. polyphemus are used to
make Limulus amebocyte lysate (LAL), which is used for the
detection of bacterial endotoxins in medical applications.[38] There is Horseshoe crab eggs
a high demand for the blood, the harvest of which involves collecting
and bleeding the animals, and then releasing them back into the sea.
Most of the animals survive the process; mortality is correlated with both the amount of blood extracted from
an individual animal, and the stress experienced during handling and transportation.[39] Estimates of mortality
rates following blood harvesting vary from 3–15%[40] to 10–30%.[41][42][43] Approximately 500,000 Limulus
are harvested annually for this purpose.[44]

Bleeding may also prevent female horseshoe crabs from being able to spawn or decrease the number of eggs
they are able to lay. Up to 30% of an individual's blood is removed, according to the biomedical industry, and
the horseshoe crabs spend between one and three days away from the ocean before being returned. As long as
the gills stay moist, they can survive on land for four days.[45] Some scientists are skeptical that certain
companies return their horseshoe crabs to the ocean at all, instead suspecting them of selling the horseshoe
crabs as fishing bait.[46]

The harvesting of horseshoe crab blood in the pharmaceutical industry is in decline. In 1986, Kyushu
University researchers discovered that the same test could be achieved by using isolated Limulus clotting
factor C (rFC), an enzyme found in LAL, as by using LAL itself.[47] Jeak Ling Ding, a National University of
Singapore researcher, patented a process for manufacturing rFC; on 8 May 2003, synthetic isolated rFC made
via her patented process became available for the first time.[48] Industry at first took little interest in the new
product, however, as it was patent-encumbered, not yet approved by regulators, and sold by a single
manufacturer, Lonza Group. In 2013, however, Hyglos GmbH also began manufacturing its own rFC product.
This, combined with the acceptance of rFC by European regulators, the comparable cost between LAL and
rFC, and support from Eli Lilly and Company, which has committed to use rFC in lieu of LAL, is projected to
all but end the practice of blood harvesting from horseshoe crabs.[49]

In June 2020, it was reported that U.S. Pharmacopeia had declined to give rFC equal standing with horseshoe
crab blood. [50] Without the approval for the classification as an industry standard testing material, U.S.
companies will have to overcome the scrutiny of showing that rFC is safe and effective for their desired uses,
which may serve as a deterrent for usage of the horseshoe crab blood substitute.[51]
In December 2019, a report of the US Senate which encouraged the Food and Drug Administration to
"establish processes for evaluating alternative pyrogenicity tests and report back [to the Senate] on steps taken
to increase their use" was released;[52] PETA backed the report.[53]

Fishery

Horseshoe crabs are used as bait to fish for eels (mostly in the United States) and whelk, or conch. However,
fishing with horseshoe crab was banned indefinitely in New Jersey in 2008 with a moratorium on harvesting
to protect the red knot, a shorebird which eats the crab's eggs.[54] A moratorium was restricted to male crabs in
Delaware, and a permanent moratorium is in effect in South Carolina.[55] The eggs are eaten in parts of
Southeast Asia, Johor and China.[56]

A low horseshoe crab population in the Delaware Bay is hypothesized to endanger the future of the red knot.
Red knots, long-distance migratory shorebirds, feed on the protein-rich eggs during their stopovers on the
beaches of New Jersey and Delaware.[57] An effort is ongoing to develop adaptive-management plans to
regulate horseshoe crab harvests in the bay in a way that protects migrating shorebirds.[58]

Shoreline development

Development along shorelines is dangerous to horseshoe crab spawning, limiting available space and
degrading habitat. Bulkheads can block access to intertidal spawning regions as well.[59]

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Further reading
Chisholm, Hugh, ed. (1911). "King-Crab". Encyclopædia Britannica (11th ed.). Cambridge
University Press.

External links
LAL Update (http://www.acciusa.com/bet/update/index.html)
Science Friday Video: horseshoe crab season (https://web.archive.org/web/20100721002111/
http://www.sciencefriday.com/program/archives/201007164)
Horseshoe crab at the Smithsonian Ocean Portal (http://ocean.si.edu/blog/flu-season-thank-hor
seshoe-crab)
The Horseshoe Crab – Medical Uses; The Ecological Research & Development Group
(ERDG) (http://www.horseshoecrab.org/med/med.html)
RedKnot.org (http://redknot.org/) links to shorebird recovery sites, movies, events & other info
on Red Knot rufa & horseshoe crabs.
Crab Bleeders (http://thefreaky.net/freaky-and-weird/crab-bleeders/) Article about the men who
bleed horseshoe crabs for science.
Day time mating of horseshoe crabs in Maine (http://naturedocumentaries.org/96/horseshoe-cra
bs-mating-Maine/)
Sarah Zhang, The Last Days of the Blue-Blood Harvest, The Atlantic, May 9, 2018 (https://ww
w.theatlantic.com/science/archive/2018/05/blood-in-the-water/559229/)

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