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The mangrove horseshoe crab (''Carcinoscorpius rotundicauda''), also known as the round-tailed horseshoe crab, is a
chelicerate The subphylum Chelicerata (from New Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. It contains the sea spiders, horseshoe crabs, and arachnids (including harvestmen, scorpions, spiders, solifuges, ticks, and mite ...
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
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 estu ...
waters in
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
,
Bangladesh Bangladesh (}, ), officially the People's Republic of Bangladesh, is a country in South Asia. It is the eighth-most populous country in the world, with a population exceeding 165 million people in an area of . Bangladesh is among the mos ...
,
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
,
Malaysia Malaysia ( ; ) is a country in Southeast Asia. The federation, federal constitutional monarchy consists of States and federal territories of Malaysia, thirteen states and three federal territories, separated by the South China Sea into two r ...
,
Brunei Brunei ( , ), formally Brunei Darussalam ( ms, Negara Brunei Darussalam, Jawi: , ), is a country located on the north coast of the island of Borneo in Southeast Asia. Apart from its South China Sea coast, it is completely surrounded by t ...
,
Singapore Singapore (), officially the Republic of Singapore, is a sovereign island country and city-state in maritime Southeast Asia. It lies about one degree of latitude () north of the equator, off the southern tip of the Malay Peninsula, bor ...
,
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is b ...
,
Cambodia Cambodia (; also Kampuchea ; km, កម្ពុជា, UNGEGN: ), officially the Kingdom of Cambodia, is a country located in the southern portion of the Indochinese Peninsula in Southeast Asia, spanning an area of , bordered by Thailan ...
,
Vietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
, China and
Hong Kong Hong Kong ( (US) or (UK); , ), officially the Hong Kong Special Administrative Region of the People's Republic of China (abbr. Hong Kong SAR or HKSAR), is a city and special administrative region of China on the eastern Pearl River Delta i ...
. It may also occur in Sri Lanka,
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John Wells explai ...
and the
Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...
, but confirmed records are lacking. Despite their name,
horseshoe crab Horseshoe crabs are marine and brackish water arthropods of the family Limulidae and the only living members of the order Xiphosura. Despite their name, they are not true crabs or crustaceans: they are chelicerates, most closely related to ar ...
s are more closely related to
spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s and
scorpion Scorpions are predatory arachnids of the order Scorpiones. They have eight legs, and are easily recognized by a pair of grasping pincers and a narrow, segmented tail, often carried in a characteristic forward curve over the back and always en ...
s (all are in the subphylum Chelicerata) than to crabs. Recent phylogenetic analysis suggests that horseshoe crabs may themselves be arachnids. The mangrove horseshoe crab is the only species in the genus ''Carcinoscorpius''. There are four extant (living) species of horseshoe crab. 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 Ko ...
'', 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 marine and brackish chelicerate arthropod. Despite their name, horseshoe crabs are more closely related to spiders, ticks, and scorpi ...
''. 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 posterior-most division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment of the arthropod body, or an additional division that is not a true segment on accou ...
) 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 an extant taxon that cosmetically 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 origin of the extant clade. Living foss ...
because they have remained practically unchanged in terms of shape and size for millions of years. Although their
physiology Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemical ...
has changed over the years, their typical three piece
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
, 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 to ...
, and
telson The telson () is the posterior-most division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment of the arthropod body, or an additional division that is not a true segment on accou ...
, has remained since the mid-
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ' ...
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 to ...
and the tail (
telson The telson () is the posterior-most division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment of the arthropod body, or an additional division that is not a true segment on accou ...
). The prosoma is the large, dome-shaped frontal part at the carapace. 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 ( ms, Semenanjung Malaysia; Jawi: سمننجڠ مليسيا), or the States of Malaya ( ms, Negeri-negeri Tanah Melayu; Jawi: نڬري-نڬري تانه ملايو), also known as West Malaysia or the Malaysian Peninsula, ...
, 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: ''Poshchim Bongo'', , abbr. WB) is a state in the eastern portion of India. It is situated along the Bay of Bengal, along with a population of over 91 million inhabitants within an area of . West Bengal is the fou ...
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 ...
and
Belawan 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 constitutes the most northerly of the city of Medan's ...
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 New Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. It contains the sea spiders, horseshoe crabs, and arachnids (including harvestmen, scorpions, spiders, solifuges, ticks, and mite ...
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 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 fangs, or similarl ...
, 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 second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") an ...
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 gills. 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 in coastal saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evolution in severa ...
. 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
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 primar ...
in the western waters, influenced by the
Pearl River The Pearl River, also known by its Chinese name Zhujiang or Zhu Jiang in Mandarin pinyin or Chu Kiang and formerly often known as the , is an extensive river system in southern China. The name "Pearl River" is also often used as a catch-a ...
. They also found that ''T. tridentatus'' can coexist in the same habitat as the mangrove horseshoe crab.


Diet

Mangrove horseshoe crabs are selective benthic feeders, feeding mainly on insect
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e, small
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of ...
,
oligochaete Oligochaeta () is a subclass of 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 megadrile earthworm ...
s, small crabs and thin-shelled bivalves. Lacking
jaw The jaw is any opposable articulated structure 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 and serv ...
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 (pterosaurs, crocodiles, alligators, dinosaurs, birds), earthworms, some gastropods, so ...
. 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 c ...
of the full and new moons. 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 The sex ratio (or gender ratio) is usually defined as 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. Many species d ...
s, such as the mangrove horseshoe crab, are found to be monogamous. In addition, the female does not choose her mate. Males find their female mates with the use of visual and chemoreceptive 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 the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
, and it takes the juveniles about five molts to grow from to adult size.


Use by humans

Thousands of the horseshoe crabs are caught by local fishermen. 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 cooked in ...
is prized as a delicacy, in Thailand most commonly served as a
salad A salad is a dish consisting of mixed, mostly natural ingredients with at least one raw ingredient. They are typically served at room temperature or chilled, though some can be served warm. Condiments and salad dressings, which exist in a va ...
called ''yam khai maeng da'' (ยำไข่แมงดา). There have occasionally been cases of food poisonings or even deaths after consuming the crabs as they were misidentified as another 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 ...
.'' Unlike larger ''T. gigas'', ''C. rotundicauda'' is known to often contain lethal
tetrodotoxin Tetrodotoxin (TTX) is a potent neurotoxin. Its name derives from Tetraodontiformes, an order that includes pufferfish, porcupinefish, ocean sunfish, and triggerfish; several of these species carry the toxin. Although tetrodotoxin was discovere ...
. In addition, horseshoe crabs are prized for their blue blood, as it is widely used in biomedical sciences for the development of drugs for diseases like mental exhaustion and gastroenteritis. The blood contains a chemical called ''Limulus'' amoebocyte lysate (LAL) that can be used to detect germs and their
endotoxins Lipopolysaccharides (LPS) are large molecules consisting of a lipid and a polysaccharide that are bacterial toxins. They are composed of an O-antigen, an outer core, and an inner core all joined by a covalent bond, and are found in the outer me ...
.


References


External links

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