Carcinoscorpius
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The mangrove horseshoe crab (''Carcinoscorpius rotundicauda''), also known as the round-tailed horseshoe crab, is a species of
horseshoe crab Horseshoe crabs are arthropods of the family Limulidae and the only surviving xiphosurans. Despite their name, they are not true crabs or even crustaceans; they are chelicerates, more closely related to arachnids like spiders, ticks, and scor ...
, a
chelicerate The subphylum Chelicerata (from Neo-Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. Chelicerates include the sea spiders, horseshoe crabs, and arachnids (including harvestmen, scorpions, spiders, solifuges, ticks ...
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
found in tropical marine and
brackish Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuari ...
waters of
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
,
Bangladesh Bangladesh, officially the People's Republic of Bangladesh, is a country in South Asia. It is the List of countries and dependencies by population, eighth-most populous country in the world and among the List of countries and dependencies by ...
, and
Southeast Asia Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...
. It may also occur in
Sri Lanka Sri Lanka, officially the Democratic Socialist Republic of Sri Lanka, also known historically as Ceylon, is an island country in South Asia. It lies in the Indian Ocean, southwest of the Bay of Bengal, separated from the Indian subcontinent, ...
,
Myanmar Myanmar, officially the Republic of the Union of Myanmar; and also referred to as Burma (the official English name until 1989), is a country in northwest Southeast Asia. It is the largest country by area in Mainland Southeast Asia and has ...
and the
Philippines The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
, but confirmed records are lacking. It is the only species in the genus ''Carcinoscorpius''. The biology, ecology and breeding patterns of ''C. rotundicauda'' and the two other Asian horseshoe crab species, ''
Tachypleus gigas ''Tachypleus gigas'', commonly known as the Indo-Pacific horseshoe crab, Indonesian horseshoe crab, Indian horseshoe crab, or southern horseshoe crab, is one of the four extant (living) species of horseshoe crab. It is found in coastal water in S ...
'' and ''
Tachypleus tridentatus ''Tachypleus tridentatus'', commonly known as the Chinese horseshoe crab, Japanese horseshoe crab, or tri-spine horseshoe crab, is a species of horseshoe crab found in Southeast and East Asia, with records from China, Indonesia, Japan, South Kore ...
'', have not been as well documented as those of the North American species ''
Limulus polyphemus The Atlantic horseshoe crab (''Limulus polyphemus''), also known as the American horseshoe crab, is a species of horseshoe crab, a kind of marine and brackish chelicerate arthropod. It is found in the Gulf of Mexico and along the Atlantic coast ...
''. All four extant species of horseshoe crabs are anatomically very similar, but ''C. rotundicauda'' is considerably smaller than the others and the only species where the cross section of the tail (
telson The telson () is the hindmost division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment (biology), segment of the arthropod body, or an additional division that is not a true segm ...
) is rounded instead of essentially triangular.


Evolutionary history

Horseshoe crabs are commonly known by biologists around the world as a
living fossil A living fossil is a Deprecation, deprecated term for an extant taxon that phenotypically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of or ...
because they have remained practically unchanged in terms of shape and size for millions of years. Although their
physiology Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
has changed over the years, their typical three piece
exoskeleton An exoskeleton () . is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton (e.g. human skeleton, that ...
, consisting of a
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 ...
,
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 ...
, and
telson The telson () is the hindmost division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment (biology), segment of the arthropod body, or an additional division that is not a true segm ...
, has remained since the mid-
Paleozoic The Paleozoic ( , , ; or Palaeozoic) Era is the first of three Era (geology), geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic (the last era of the Proterozoic Eon) and ends 251.9 Ma a ...
era. Fossils of horseshoe crabs that have been dated to over look almost identical to those species that are still alive today. The long existence of this body plan suggests its success. The three Asian species of horseshoe crabs form their own clade, and it is estimated that they diverged from the American species between to .


Anatomy

The basic body plan of a horseshoe crab consists of three parts: the
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 ...
, the
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 ...
and the tail (
telson The telson () is the hindmost division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment (biology), segment of the arthropod body, or an additional division that is not a true segm ...
). The prosoma is the large, dome-shaped frontal part at the
carapace A carapace is a 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 tortoises, the unde ...
. The smaller rear carapace with spines on the edge is the opisthosoma. The rear extension that looks like a spike is the telson, which is commonly described as the tail. Uniquely among the horseshoe crabs, the cross section of the tail of the mangrove horseshoe crab is rounded. It is essentially triangular in the other species. The tail is used to turn itself right side up when overturned. The mangrove horseshoe crab is the smallest of the four living species of horseshoe crabs. Like the other species, females grow larger than males. On average in
Peninsular Malaysia Peninsular Malaysia, historically known as Malaya and also known as West Malaysia or the Malaysian Peninsula, is the western part of Malaysia that comprises the southern part of the Malay Peninsula on Mainland Southeast Asia and the list of isla ...
, females are about long, including a tail that is about , and their carapace (prosoma) is about wide. In comparison, the average for males is about long, including a tail that is about , and their carapace is about wide. There are significant geographic variations in the size, but this does not follow a clear north–south or east–west pattern. Those from
West Bengal West Bengal (; Bengali language, Bengali: , , abbr. WB) is a States and union territories of India, state in the East India, eastern portion of India. It is situated along the Bay of Bengal, along with a population of over 91 million inhabi ...
in India average somewhat smaller than those from Peninsular Malaysia, with a carapace width of about and in females and males respectively. Elsewhere it averages even smaller, with the smallest reported from the
Balikpapan Balikpapan is a seaport city in East Kalimantan, Indonesia. Located on the east coast of the island of Borneo, the city is the financial center of Kalimantan. Balikpapan is the city with the largest economy in Kalimantan with an estimated 20 ...
and
Belawan The Port of Belawan ( zh, t=勿老灣, s=, poj=mài lau ôan) is a harbor in Medan, North Sumatra. Located on the northeast coast of Sumatra, Belawan is Indonesia's busiest seaport outside of Java. It includes Medan Belawan District, the most nor ...
regions in Indonesia where the carapace width of females is about and in males . The largest females of the species may reach up to in length, including the tail. Uncommon for
chelicerate The subphylum Chelicerata (from Neo-Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. Chelicerates include the sea spiders, horseshoe crabs, and arachnids (including harvestmen, scorpions, spiders, solifuges, ticks ...
s, horseshoe crabs have two
compound eyes A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which distin ...
. The main function of these compound eyes is to find a mate. In addition, they have two median eyes, two rudimentary lateral eyes, and an endoparietal eye on their carapace and two ventral eyes located on the underside by the mouth. Scientists believe the two ventral eyes aid in the orientation of the horseshoe crab when swimming. Each individual has six pairs of appendages. The first pair, the
chelicerae The chelicerae () are the arthropod mouthparts, mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as "jaws", chelicerae may be shaped as either articulated ...
, are relatively small and placed in front of the mouth. They are used to place food in it. The remaining five pairs of legs are placed on either side of the mouth and are used for walking/pushing. These are the
pedipalp Pedipalps (commonly shortened to palps or palpi) are the secondary pair of forward appendages among Chelicerata, chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to ...
s (first pair) and the pusher legs (remaining four pairs). Most of the appendages have straight, scissor-like claws, but in males the first and second pair of walking legs have strongly hooked "scissors", which are used for grasping the female during mating. Located behind their legs are
book gill A book is a structured presentation of recorded information, primarily verbal and graphical, through a medium. Originally physical, electronic books and audiobooks are now existent. Physical books are objects that contain printed material, mo ...
s. These gills are used for propulsion to swim and to exchange respiratory gases.


Distribution and habitat

This species occurs only in Asia around the Indo-West Pacific region where the climate is tropical or subtropical. These horseshoe crabs can be found to exist throughout southeast Asia in shallow waters with soft, sandy bottoms or extensive mud flats. The mangrove horseshoe crab is benthopelagic, spending most of its life close to or at the bottom of a body of their brackish, swampy water habitat, such as
mangroves A mangrove is a shrub or tree that grows mainly in coastal saline or brackish water. Mangroves grow in an equatorial climate, typically along coastlines and tidal rivers. They have particular adaptations to take in extra oxygen and remove sal ...
. This is the habitat for which it gains its common name: mangrove horseshoe crab. Scientists have studied the distribution of mangrove horseshoe crabs in Hong Kong specifically. The researchers noted their abundance on the beaches of Hong Kong before the sharp decline in population over the past ten years. In the study, they found an uneven distribution of the horseshoe crabs throughout Hong Kong, with a greater abundance found in the western waters. They predict this unevenness is due to the
estuarine An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environm ...
hydrography Hydrography is the branch of applied sciences which deals with the measurement and description of the physical features of oceans, seas, coastal areas, lakes and rivers, as well as with the prediction of their change over time, for the primary ...
in the western waters, influenced by the
Pearl River The Pearl River (, or ) is an extensive river system in southern China. "Pearl River" is often also used as a catch-all for the watersheds of the Pearl tributaries within Guangdong, specifically the Xi ('west'), Bei ('north'), and Dong ( ...
. They also found that ''T. tridentatus'' can coexist in the same habitat as the mangrove horseshoe crab.


Diet

Mangrove horseshoe crabs are selective
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
feeders, feeding mainly on insect
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e, small
fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
,
oligochaete Oligochaeta () is a subclass of soft-bodied animals in the phylum Annelida, which is made up of many types of aquatic and terrestrial worms, including all of the various earthworms. Specifically, oligochaetes comprise the terrestrial megadril ...
s, small
crab Crabs are decapod crustaceans of the infraorder Brachyura (meaning "short tailed" in Greek language, Greek), which typically have a very short projecting tail-like abdomen#Arthropoda, abdomen, usually hidden entirely under the Thorax (arthropo ...
s and thin-shelled
bivalves Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed by a calcified exoskeleton consis ...
. Lacking
jaw The jaws are a pair of opposable articulated structures at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth ...
s, it grinds up the food with bristles on its legs and places it in its mouth using its chelicerae. The ingested food then enters the cuticle-lined oesophagus and then the
proventriculus The proventriculus is part of the digestive system of birds.Encarta World English Dictionary orth American Edition(2007). ''Proventriculus''. Source: (accessed: December 18, 2007) An analogous organ exists in invertebrates and insects. Birds Th ...
. The proventriculus is made up of a crop and a
gizzard The gizzard, also referred to as the ventriculus, gastric mill, and gigerium, is an organ found in the digestive tract of some animals, including archosaurs (birds and other dinosaurs, crocodiles, alligators, pterosaurs), earthworms, some gast ...
. The crop can expand to fit the ingested food, while the gizzard grinds the food into a pulp. Studies have found that mangrove horseshoe crabs have a strong preference for insect larvae over the other organisms on which it also feeds.


Mating biology

In the spring, horseshoe crabs migrate from the deeper water to the shallow, muddy areas. Nesting usually follows the cycle of the
high tide Tides are the rise and fall of sea levels caused by the combined effects of the gravitational forces exerted by the Moon (and to a much lesser extent, the Sun) and are also caused by the Earth and Moon orbiting one another. Tide tables ...
of the full and
new moon In astronomy, the new moon is the first lunar phase, when the Moon and Sun have the same ecliptic longitude. At this phase, the lunar disk is not visible to the naked eye, except when it is silhouetted against the Sun during a solar eclipse. ...
s. During the mating period, the males will follow and cling to the backs of their potential mates using modified prosomal appendages for long periods of time before the egg-laying has occurred. Horseshoe crab species with low spawning densities and 1:1
sex ratio A sex ratio is the ratio of males to females in a population. As explained by Fisher's principle, for evolutionary reasons this is typically about 1:1 in species which reproduce sexually. However, many species deviate from an even sex ratio, ei ...
s, such as the mangrove horseshoe crab, are found to be
monogamous Monogamy ( ) is a relationship of two individuals in which they form a mutual and exclusive intimate partnership. Having only one partner at any one time, whether for life or serial monogamy, contrasts with various forms of non-monogamy (e.g. ...
. In addition, the female does not choose her mate. Males find their female mates with the use of visual and
chemoreceptive A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...
signals. Once a mate is found, the female digs a hole and lays the eggs while the male externally fertilizes them. Once the eggs are laid, the male and female go back to the ocean, and the eggs develop on their own. Their eggs are large, and after a couple weeks, the eggs hatch into miniature versions of the adults. The females lay about 3500 of them. Mangrove horseshoe crabs in Singapore breed from August to April. Juveniles grow about 33% bigger each time they
molt In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is a process by which an animal casts off parts of its body to serve some beneficial purpose, either at ...
, and it takes the juveniles about five molts to grow from to adult size.


Use by humans

Mangrove horseshoe crabs are collected by fishermen throughout Southeast Asia for culinary use. While the crabs have very little flesh, their
roe Roe, ( ) or hard roe, is the fully ripe internal egg masses in the ovaries, or the released external egg masses, of fish and certain marine animals such as shrimp, scallop, sea urchins and squid. As a seafood, roe is used both as a cooking, c ...
is prized as a delicacy, in Thailand most commonly served as a
salad A salad is a dish consisting of mixed ingredients, frequently vegetables. They are typically served chilled or at room temperature, though some can be served warm. Condiments called '' salad dressings'', which exist in a variety of flavors, a ...
called ''yam khai maeng da'' (ยำไข่แมงดา). Unlike ''
Tachypleus gigas ''Tachypleus gigas'', commonly known as the Indo-Pacific horseshoe crab, Indonesian horseshoe crab, Indian horseshoe crab, or southern horseshoe crab, is one of the four extant (living) species of horseshoe crab. It is found in coastal water in S ...
'', the flesh of ''C. rotundicauda'' is known to contain lethal
tetrodotoxin Tetrodotoxin (TTX) is a potent neurotoxin. Its name derives from Tetraodontiformes, an Order (biology), order that includes Tetraodontidae, pufferfish, porcupinefish, ocean sunfish, and triggerfish; several of these species carry the toxin. Alt ...
(TTX). There have been several cases of food poisoning, some resulting in death, after consuming mangrove horseshoe crab. Typically, this is due to being misidentified as ''T. gigas'', which is harmless to consume.


References


External links

* {{Taxonbar, from=Q2673503 Xiphosura Arthropods of China Fauna of Hong Kong Animals described in 1985 Taxonomy articles created by Polbot